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Species are altering their ranges as a response to climate change, but the magnitude and direction of observed range shifts vary considerably among species. The ability to persist in current areas and colonize new areas plays a crucial role in determining which species will thrive and which decline as climate change progresses. Several studies have sought to identify characteristics, such as morphological and life-history traits, that could explain differences in the capability of species to shift their ranges together with a changing climate. These characteristics have explained variation in range shifts only sporadically, thus offering an uncertain tool for discerning responses among species. As long-term selection to past climates have shaped species’ tolerances, metrics describing species’ contemporary climatic niches may provide an alternative means for understanding responses to on-going climate change. Species that occur in a broader range of climatic conditions may hold greater tolerance to climatic variability and could therefore more readily maintain their historical ranges, while species with more narrow tolerances may only persist if they are able to shift in space to track their climatic niche. Here, we provide a first-filter test of the effect of climatic niche dimensions on shifts in the leading range edges in three relatively well-dispersing species groups. Based on the realized changes in the northern range edges of 383 moth, butterfly, and bird species across a boreal 1,100 km latitudinal gradient over c. 20 years, we show that while most morphological or life-history traits were not strongly connected with range shifts, moths and birds occupying a narrower thermal niche and butterflies occupying a broader moisture niche across their European distribution show stronger shifts towards the north. Our results indicate that the climatic niche may be important for predicting responses under climate change and as such warrants further investigation of potential mechanistic underpinnings.

Aapa mires are EU priority habitats that harbour unique biodiversity values but face increasing global change threats. Here, we investigate the exposure of red-listed aapa mire species inhabiting fen and flark fen habitats to the impacts of land use and climate change. Climate change-based threats were assessed across the aapa mire zone of Finland based on climate velocities (a metric describing the speed and direction of climate movement) measured for mean January temperature (TJan), growing degree days (GDD5) and mean annual water balance (WAB). Land use threats were assessed based on the cover of drainage ditches and three other adverse land use types around the species occurrences. Our results suggest that rapid changes in TJan may alter winter thermal conditions and thereby also species performance, particularly in the northernmost part of the aapa mire zone, where the most valuable concentrations of red-listed species are situated. The land use and GDD5 threats are highest in the southern regions where the red-listed aapa mire species occurrences are sparser but face severe risks to their persistence. In the central part of the aapa mire zone, a number of valuable aapa mires with red-listed species are exposed to both intermediately high TJan and GDD5 velocities and a spatially varying amount of ditching. Three conservation approaches to support the persistence of red-listed aapa mire species: (i) restora-tion, (ii) establishment of new protected areas, and (iii) monitoring of the key habitats, should be flexibly and complementarily applied to the preservation of aapa mires subject to accelerating climate change.

Harsh winters are a characteristic element of Arctic ecosystems, yet the importance of winter conditions for Arctic plant communities is still underrepresented in climate change impact studies. Here, we use fine-scale microclimate data with plot-scale records of vascular plants, lichens and bryophytes from three Arctic areas, and show that topographically heterogeneous tundra holds marked spatial variation, especially in winter temperatures. Winter conditions are the strongest environmental variable related to the fine-scale patterns in tundra vegetation, whereas summer temperatures mainly explain the coarse-scale differences among the Arctic areas. Nonetheless, the three plant groups (and also individual species) show often contrasting and complex responses along the local environmental gradients. Our results highlight the importance of local conditions and heterogeneity for tundra plants, and knowing that the Arctic winters are warming faster than summers, a greater focus should be placed on winter conditions in simulations of climate change impacts in tundra ecosystems. Winter conditions are the strongest environmental variable relating to fine-scale patterns in tundra vegetation. Vascular plants, lichens and bryophytes nonetheless show complex responses across environmental gradients, highlighting the importance of local heterogeneity.

Publisher Copyright: © 2021 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America. The use of indicator species in forest conservation and management planning can facilitate enhanced preservation of biodiversity from the negative effects of forestry and other uses of land. However, this requires detailed and spatially comprehensive knowledge of the habitat preferences and distributions of selected focal indicator species. Unfortunately, due to limited resources for field surveys, only a small proportion of the occurrences of focal species is usually known. This shortcoming can be circumvented by using modelling techniques to predict the spatial distribution of suitable sites for the target species. Airborne laser scanning (ALS) and other remote sensing (RS) techniques have the potential to provide useful environmental data covering systematically large areas for these purposes. Here, we focused on six bird of prey and woodpecker species known to be good indicators of boreal forest biodiversity values. We used known nest sites of the six indicator species based on nestling ringing records. Thus, the most suitable nesting sites of these species provide important information for biodiversity-friendly forest management and conservation planning. We developed fine-grained, i.e., 96 x 96 m grid cell resolution, predictive maps across the whole of Finland of the suitable nesting habitats based on ALS and other RS data and spatial information on the distribution of important forest stands for the six studied biodiversity indicator bird species based on nesting habitat suitability modelling, i.e., the MaxEnt model. Habitat preferences of the study species, as determined by MaxEnt, were in line with the previous knowledge of species-habitat relations. The proportion of suitable habitats of these species in protected areas was considerable, but our analysis also revealed many potentially high-quality forest stands outside protected areas. However, many of these sites are increasingly threatened by logging due to increased pressures for using forests for bioeconomy and forest industry based on National Forest Strategy. Predicting habitat suitability based on information on the nest sites of indicator species provides a new tool for systematic conservation planning over large areas in boreal forests in Europe, and corresponding approach would also be feasible and recommendable elsewhere where similar data are available. The use of indicator species in forest conservation and management planning can facilitate enhanced preservation of biodiversity from the negative effects of forestry and other uses of land. However, this requires detailed and spatially comprehensive knowledge of the habitat preferences and distributions of selected focal indicator species. Unfortunately, due to limited resources for field surveys, only a small proportion of the occurrences of focal species is usually known. This shortcoming can be circumvented by using modeling techniques to predict the spatial distribution of suitable sites for the target species. Airborne laser scanning (ALS) and other remote sensing (RS) techniques have the potential to provide useful environmental data covering systematically large areas for these purposes. Here, we focused on six bird of prey and woodpecker species known to be good indicators of boreal forest biodiversity values. We used known nest sites of the six indicator species based on nestling ringing records. Thus, the most suitable nesting sites of these species provide important information for biodiversity-friendly forest management and conservation planning. We developed fine-grained, that is, 96 x 96 m grid cell resolution, predictive maps across the whole of Finland of the suitable nesting habitats based on ALS and other RS data and spatial information on the distribution of important forest stands for the six studied biodiversity indicator bird species based on nesting-habitat suitability modeling, that is, the MaxEnt model. Habitat preferences of the study species, as determined by MaxEnt, were in line with the previous knowledge of species-habitat relations. The proportion of suitable habitats of these species in protected areas (PAs) was considerable, but our analysis also revealed many potentially high-quality forest stands outside PAs. However, many of these sites are increasingly threatened by logging because of increased pressures for using forests for bioeconomy and forest industry based on National Forest Strategy. Predicting habitat suitability based on information on the nest sites of indicator species provides a new tool for systematic conservation planning over large areas in boreal forests in Europe, and a corresponding approach would also be feasible and recommendable elsewhere where similar data are available.

In this paper, we present daily gridded evapotranspiration dataset covering Finland over the 40-year period 1981–2020. The dataset representing evapotranspiration from a reference surface is freely available in a 1 km × 1 km grid.

Climate velocity is an increasingly used metric to detect habitats, locations and regions which are exposed to high rates of climate change and displacement. In general, velocities are measured based on the assumption that future climatically similar locations can occur anywhere in the study landscape. However, this assumption can provide a biased basis for habitats which are constrained to specific environmental conditions. For such habitats, a set of selected suitable locations may provide ecologically more realistic velocity measures. Here, we focus on one environmentally constrained habitat, aapa mires, which are peat-accumulating EU Habitats Directive priority habitats, whose ecological conditions and biodiversity values may be jeopardised by climate change. We assess the climate exposure of aapa mires in Finland by developing velocity metrics separately for the whole ≥10 ha aapa mire complexes (‘aapa mires’) and their wettest flark-dominated parts (‘flark fens’). Velocity metrics were developed for six bioclimatic variables (growing degree days (GDD5), mean January and July temperatures, annual precipitation, and May and July water balance, based on climate data for 1981–2010 and for 2040–2069 as derived from global climate models for two Representative Concentration Pathways (RCP4.5 and RCP8.5). For the six variables, velocities were calculated based on the distance between climatically similar present-day and nearest future mire, divided by the number of years between the two periods, and by excluding the unsuitable matrix. Both aapa mires and flark fens showed high exposure (>5 km/year) to changes in January temperature, and often also considerably high velocities for GDD5 and July temperatures. The flark fens showed significantly higher climate velocities than the aapa mires and had a smaller amount of corresponding habitat in their surroundings. Thus, many of the studied mires, particularly the flark fens, are likely to face increased risks of exposure due to changes in winter and summer temperatures. Moreover, considerable changes in precipitation-related conditions may occur at the southern margin of the aapa mire zone. Our results show that specifically tailored climate velocity metrics can provide a useful quantitative tool to inform conservation and management decisions to support the ecosystem sustainability of this EU Habitats Directive biotope and targeting restoration towards the most vulnerable aapa mires. Highlights • Aapa mires are EU Habitats Directive priority habitats of high conservation value • Velocity assessments enable quantifying the exposure of habitats to climate change • Boreal aapa mires will face notable exposure risks due to warming climate • Wet flark fens with open water pools and their biodiversity are particularly at risk • Climate velocity information supports adaptive biodiversity conservation planning

Recent actions to mitigate biodiversity loss in agricultural environments appear insufficient despite the considerable efforts channeled via the European Union’s Common Agricultural Policy. One likely reason for this failure is the limited attention paid to the regional and landscape level ecological characteristics in farmland conservation planning. We demonstrate how to obtain conservation prioritization solutions that would address simultaneously three goals, including two landscape level targets: minimizing local habitat quality loss, maximizing habitat connectivity, and incorporating landscape heterogeneity. As these goals may be contradictory, we investigate the potential trade-offs between them. We used the Zonation prioritization tool to examine how our three goals could be implemented in the agricultural landscapes of southwest Finland. We used measures of (i) biodiversity value of grasslands, (ii) connectivity between grasslands, and (iii) landscape heterogeneity which comprised of (land cover type based) compositional heterogeneity and (field margin based) configurational heterogeneity. Integration of landscape heterogeneity measures and habitat connectivity resulted in some tradeoffs with local habitat quality, the most prominent observation being that landscape heterogeneity co-varied with grassland connectivity. Among the two landscape heterogeneity parameters, inclusion of compositional heterogeneity resulted in more clustered prioritization solutions than configurational heterogeneity, which had a spatially more balanced impact. Concordance among landscape scale factors implies high potential for reconstruction of a functioning network of semi-natural grasslands in areas under intensive agricultural use. Broader scale multi-objective planning approaches can thus importantly support targeting biodiversity conservation planning and mediating the implementation of Common Agricultural Policy objectives. peerReviewed

Biodiversity crisis calls for better knowledge of the status of biodiversity, but also drivers and pressures behind the ecosystem change and the vulnerability of habitats. There is growing demand for the biodiversity data for decision-making on national, regional and global scales. However, the operationalized biodiversity and ecosystem monitoring systems which can integrate remote sensing, in situ and modelling data are still rare despite of that there are good co-operation initiatives such as Australian TERN and US NEON in place. In Finland, we have launched the Finnish Ecosystem Observatory (FEO) as a reaction to such a need to upgrade and modernize biodiversity and ecosystem monitoring schemes. We have started a development of national research and monitoring infrastructure/platform which can integrate different data types and provide timely data and knowledge for various decision-making purposes. In this short paper, we describe three case studies as examples of different data types and different monitoring needs: i) habitat mapping for large area, ii) hydrological condition of aapa mires, and iii) automated Aspen mapping for boreal forests. We will evaluate the possibilities to operationalize the production of such data sets in a long-term as well as the challenges and bottlenecks related either to data, processing or applicability. Finally, we will also discuss how FEO and similar kind of biodiversity monitoring platforms could be further developed, and how the network of such observatories could co-operate in the future.

Highlights • Climate change-induced risks to EU Habitats Directive species are poorly known. • Adverse land use for species often coincides with high climate change velocity. • Climate change and land use threats vary notably both between and within species. • Mapping risks across species occurrences is central to climate-wise conservation. • Climate change land-use risks hinder Habitats Directive species from reaching a favourable status. The Habitats Directive of the European Union is a key legislative instrument in Europe, supporting the conservation of rare, threatened or endemic species. It aims at ensuring that the species listed in the Annexes of the Habitats Directive show a favourable conservation status, i.e., that they are able to maintain viable populations and that their natural range is sufficient and not decreasing currently, nor will in the future. However, climate change may hamper Habitats Directive species in achieving (or maintaining) a favourable conservation status, particularly when these impacts are amplified by adverse land use. Here, we studied Habitats Directive species in Finland for which ≥70% of the occurrences were recorded with the resolution of ≤100 m. The number of occurrence sites for the 52 species studied ranged from one site to 13,653 sites, summing up to 19,367 sites. For all these sites and their surroundings, we assessed the vulnerabilities caused by climate change and land use. The climate exposure of occurrence sites was measured based on the rapidity of climatic changes (i.e. climate velocity) in three climate variables (growing degree days, mean January air temperature, water balance) at each site. Risks caused by land use were assessed using two negative and four positive variables that respectively described the quantity of land cover and habitats that is either harmful (e.g. clear-cut forest and drained peatlands) or supportive (protected areas and suitable habitats) to species occurrences. To complement climate and land-use variables, three additional variables describing protection status of the sites and the number of occurrences of the same species in the landscape were examined. Comparison of the mean vulnerability values for each species showed that some of the species inhabit, on average, areas with high climate exposure. Moreover, in certain species climate change-induced vulnerabilities consistently coincide with negative land use. However, in many of the 52 species there was large variation in the vulnerability levels between individual occurrence sites, concerning both climate exposure and land-use variables. Considering the vulnerabilities due to climate change separately, 40–60% of the species occurrence sites are expected to face high exposure caused by rapid changes in summer or winter temperatures, which presents challenges in maintaining a favourable conservation status. Our results also revealed numerous species occurrences where high climate velocity coincided with a large amount of negative land use and low amount of suitable habitat, for which climate-wise conservation planning could be targeted.

The challenges posed by climate change and biodiversity loss are deeply interconnected. Successful co-managing of these tangled drivers requires innovative methods that can prioritize and target management actions against multiple criteria, while also enabling cost-effective land use planning and impact scenario assessment. This paper synthesises the development and application of an integrated multidisciplinary modelling and evaluation framework for carbon and biodiversity in forest systems. By analysing and spatio-temporally modelling carbon processes and biodiversity elements, we determine an optimal solution for their co-management in the study landscape. We also describe how advanced Earth Observation measurements can be used to enhance mapping and monitoring of biodiversity and ecosystem processes. The scenarios used for the dynamic models were based on official Finnish policy goals for forest management and climate change mitigation. The development and testing of the system were executed in a large region in southern Finland (Kokemäenjoki basin, 27 024 km2) containing highly instrumented LTER (Long-Term Ecosystem Research) stations; these LTER data sources were complemented by fieldwork, remote sensing and national data bases. In the study area, estimated total net emissions were currently 4.2 TgCO2eq a-1, but modelling of forestry measures and anthropogenic emission reductions demonstrated that it would be possible to achieve the stated policy goal of carbon neutrality by low forest harvest intensity. We show how this policy-relevant information can be further utilised for optimal allocation of set-aside forest areas for nature conservation, which would significantly contribute to preserving both biodiversity and carbon values in the region. Biodiversity gain in the area could be increased without a loss of carbon-related benefits. The challenges posed by climate change and biodiversity loss are deeply interconnected. Successful co-managing of these tangled drivers requires innovative methods that can prioritize and target management actions against multiple criteria, while also enabling cost-effective land use planning and impact scenario assessment. This paper synthesises the development and application of an integrated multidisciplinary modelling and evaluation framework for carbon and biodiversity in forest systems. By analysing and spatio-temporally modelling carbon processes and biodiversity elements, we determine an optimal solution for their co-management in the study landscape. We also describe how advanced Earth Observation measurements can be used to enhance mapping and monitoring of biodiversity and ecosystem processes. The scenarios used for the dynamic models were based on official Finnish policy goals for forest management and climate change mitigation. The development and testing of the system were executed in a large region in southern Finland (Kokemäenjoki basin, 27,024 km2) containing highly instrumented LTER (Long-Term Ecosystem Research) stations; these LTER data sources were complemented by fieldwork, remote sensing and national data bases. In the study area, estimated total net emissions were currently 4.2 TgCO2eq a−1, but modelling of forestry measures and anthropogenic emission reductions demonstrated that it would be possible to achieve the stated policy goal of carbon neutrality by low forest harvest intensity. We show how this policy-relevant information can be further utilized for optimal allocation of set-aside forest areas for nature conservation, which would significantly contribute to preserving both biodiversity and carbon values in the region. Biodiversity gain in the area could be increased without a loss of carbon-related benefits.

Intensive management of boreal forests impairs forest biodiversity and species of old-growth forest. Effective measures to support biodiversity require detection of locations valuable for conservation. We applied species distribution models (SDMs) to a species of mature forest, the northern goshawk (Accipiter gentilis, goshawk), that is often associated with hotspots of forest biodiversity. We located optimal sites for the goshawk on a landscape scale, assessed their state under intensified logging operations and identified characteristics of goshawks' nesting sites in boreal forests. Optimal sites for the goshawk covered only 3.4% of the boreal landscape and were mostly located outside protected areas, which highlights the importance of conservation actions in privately-owned forests. Furthermore, optimal sites for the goshawk and associated biodiversity were under threat. Half of them were logged to some extent and 10% were already lost or notably deteriorated due to logging shortly after 2015 for which our models were calibrated. Habitat suitability for the goshawk increased with increasing volume of Norway spruce (Picea abies) peaking at 220 m(3) ha(-1), and with small quantities of birches (Betula spp.) and other broad-leaved trees. Threats to biodiversity of mature spruce forests are likely to accelerate in the future with increasing logging pressures and shorter rotation periods. Logging should be directed less to forests with high biodiversity. Continuous supply of mature spruce forests in the landscape should be secured with a denser network of protected areas and measures that aid in sparing large entities of mature forest on privately-owned land. peerReviewed

Significance Plant functional traits are central instruments in developing understanding and predicting biodiversity patterns and ecosystems processes. Snow is an important ecological factor in cold climates, but its contribution to the evolution of functionality of tundra vegetation is poorly known and insufficiently addressed in the research. We show here that snow has a fundamental effect in mediating climate change impacts on functional composition and diversity of Arctic tundra vegetation. As a whole, Arctic landscapes may lose spatial heterogeneity because plant communities will be functionally more alike, although the local functional diversity may increase. Our results highlight that future snow conditions and their fine-scale variability should be acknowledged in the next generation of Arctic vegetation−ecosystem models., The Arctic is one of the least human-impacted parts of the world, but, in turn, tundra biome is facing the most rapid climate change on Earth. These perturbations may cause major reshuffling of Arctic species compositions and functional trait profiles and diversity, thereby affecting ecosystem processes of the whole tundra region. Earlier research has detected important drivers of the change in plant functional traits under warming climate, but studies on one key factor, snow cover, are almost totally lacking. Here we integrate plot-scale vegetation data with detailed climate and snow information using machine learning methods to model the responsiveness of tundra communities to different scenarios of warming and snow cover duration. Our results show that decreasing snow cover, together with warming temperatures, can substantially modify biotic communities and their trait compositions, with future plant communities projected to be occupied by taller plants with larger leaves and faster resource acquisition strategies. As another finding, we show that, while the local functional diversity may increase, simultaneous biotic homogenization across tundra communities is likely to occur. The manifestation of climate warming on tundra vegetation is highly dependent on the evolution of snow conditions. Given this, realistic assessments of future ecosystem functioning require acknowledging the role of snow in tundra vegetation models.

The amount of High Nature Value farmland (HNVf) is a commonly used environmental indicator for assessing the performance of the Common Agricultural Policy, to support sustainable agriculture and monitor changes in agricultural land use in Europe. HNVf comprises agricultural areas of semi-natural state, low-intensity farming and fine-scale landscape mosaics of different habitat types. For a successful implementation, the identification of HNVf should correctly reflect the variation in biodiversity values between different agricultural landscapes. We examined how well the Finnish HNVf indicator and the sub-indicators constituting it – recalculated for the purposes of this study for five study regions – reflect the variation in bird and butterfly species richness and diversity patterns at different spatial scales. We found that butterfly diversity index was positively associated with the HNVf indicator at the finest scale of 0.5 km × 0.5 km squares. Among the HNVf sub-indicators, extensive cultivation of grasslands was most strongly related to the farmland bird diversity and the density of edge to the butterfly diversity. Thus, the HNVf concept reflects well the distribution of butterflies in the Finnish agricultural landscapes but insufficiently the diversity patterns of farmland birds. Importantly, semi-natural vegetation and long-term pastures – the backbone of the concept – presently occur in small and highly fragmented patches in agricultural landscapes in Finland. The Pan-European concept of HNVf has restricted application to farmland birds of this boreal country and the national HNVf concept may need to be revised.

Climate change velocity is an increasingly used metric to assess the broad-scale climatic exposure and climate change induced risks to terrestrial and marine ecosystems. However, the utility of this metric in conservation planning can be enhanced by determining the velocities of multiple climatic drivers in real protected area (PA) networks on ecologically relevant scales. Here we investigate the velocities of three key bioclimatic variables across a nation-wide reserve network, and the consequences of including fine-grained topoclimatic data in velocity assessments. Using 50-m resolution data describing present-day and future topoclimates, we assessed the velocities of growing degree days, the mean January temperature and climatic water balance in the Natura 2000 PA network in Finland. The high-velocity areas for the three climate variables differed drastically, indicating contrasting exposure risks in different PAs. The 50-m resolution climate data revealed more realistic estimates of climate velocities and more overlap between the present-day and future climate spaces in the PAs than the 1-km resolution data. Even so, the current temperature conditions were projected to disappear from almost all the studied PAs by the end of this century. Thus, in PA networks with only moderate topographic variation, far-reaching climate change induced ecological changes may be inevitable.

Increased attention is being paid to the ecological drivers and conservation measures which could mitigate climate change-induced pressures for species survival, potentially helping populations to remain in their present-day locations longer. One important buffering mechanism against climate change may be provided by the heterogeneity in topography and consequent local climate conditions. However, the buffering capacity of this topoclimate has so far been insufficiently studied based on empirical survey data across multiple sites and species. Here, we studied whether the fine-grained air temperature variation of protected areas (PAs) affects the population changes of declining northern forest bird species. Importantly to our study, in PAs harmful land use, such as logging, is not allowed, enabling the detection of the effects of temperature buffering, even at relatively moderate levels of topographic variation. Our survey data from 129 PAs located in the boreal zone in Finland show that the density of northern forest species was higher in topographically heterogeneous PAs than in topographically more homogeneous PAs. Moreover, local temperature variation had a significant effect on the density change of northern forest birds from 1981&ndash, 1999 to 2000&ndash, 2017, indicating that change in bird density was generally smaller in PAs with higher topographic variation. Thus, we found a clear buffering effect stemming from the local temperature variation of PAs in the population trends of northern forest birds.

Snow has far-reaching effects on ecosystem processes and biodiversity in high-latitude ecosystems, but these have been poorly considered in climate change impact models1,2. Here, to forecast future trends in species occurrences and richness, we fitted species–environment models with temperature data from three climate scenarios and simulated up to a 40% decrease in snow cover duration (SCD)3. We used plot-scale data on 273 vascular plant, moss and lichen species in 1,200 study sites spanning a wide range of environmental conditions typical for mountainous Arctic landscapes (within 165 km2). According to the models, a rise in temperature increased overall species richness and caused only one species to lose all suitable habitat. In contrast, a shorter SCD tempered the effect of increasing temperature on species richness and led to accelerated rates of species’ local extinctions after a tipping point at 20–30% SCD decrease. All three species groups showed similar extinction rates but contrasting species richness responses. Our simulations indicate that future biodiversity patterns in Arctic regions are highly dependent on the evolution of snow conditions. Climate impact models that ignore the effects of snow cover change may provide biased biodiversity projections, with potentially erratic implications for Arctic nature conservation planning. Arctic biodiversity patterns will be highly dependent on the evolution of snow conditions, according to simulation results that integrate observations of vascular plants, mosses and lichens over a range of Arctic landscapes.

Climate change has the potential to have wholesale impacts on species populations, driving them polewards and upwards, and even affecting populations occurring within protected area (PA) networks. We studied population changes in bird species in the boreal PA network of Finland based on extensive bird census data collected in the years 1981–1999 and in 2000–2017. Between these time periods, the mean annual temperature increased in Finland by 1.1 °C, and the mean weighted density of the species shifted 28.5 km (1.8 km/year) northwards in the PA network. However, the total bird population density simultaneously declined by approximately 10%. The decline was most pronounced in long-distance migrants, which showed strongest population contraction in southern boreal regions. In contrast, resident species increased between the two time slices, particularly in larger PAs. While the PA network of boreal native habitats appears to be successful in preserving resident species, climate-induced changes have also caused a decline in the populations of migrants in the PAs. Thus, life-history characteristics of species can significantly affect the success of conservation efforts in a warming climate. To enhance future survival of resident and migrant bird species moving to Finland and northwards, the PAs should be larger and the connectivity of the PA network improved in southern and central Finland. In addition, international actions are needed to enhance the survival of long-distance migrants during the migratory period and in overwintering grounds.

Agricultural intensification has caused drastic declines in the area and species richness of semi-natural grasslands across Europe. Novel habitats, such as power line clearings, provide alternative habitats and niches for grassland species, and might therefore mitigate these declines. However, it is not fully understood which environmental factors determine the occurrence of grassland species in the clearings. Identifying the most important drivers for grassland species occurrence would help understand the value of the clearings for grassland conservation and target enhanced management into clearings with most potential as grassland habitat. We studied the effects of local environmental conditions, and past and present connectivity to semi-natural grasslands, on the species richness of grassland plants and butterflies in 43 power line clearings in Finland. The results of generalized linear models and hierarchical partitioning showed that increasing time since clear-cut and amount of clearing residue decreased the species richness of both species groups, while the cover of mesic habitats increased it. However, the two species groups showed also divergent responses. Present-day local environmental conditions appeared to be the sole driver of grassland butterfly species richness, whereas the richness of grassland plants was related both to current conditions and historical connectivity to grasslands in 1870–1880s. This suggests the presence of an extinction debt in the studied grassland plant communities, emphasizing the need for enhanced management to increase suitable grassland habitat in the clearings. This would diminish the potential future losses of grassland plant species in the clearings and create valuable habitat for grassland butterflies as well.

All Nordic countries are committed to implement the Convention on Biological Diversity (CBD) and the CBD Strategic Plan for Biodiversity 2011-2020. According to the CBD Aichi Targets, areas of particular importance for biodiversity and ecosystem services should be protected through ecologically representative and well-connected systems of protected areas as well as other effective area-based conservation measures. The ecosystems of the northern parts of Norway, Sweden, Finland and North-West Russia are very diverse and unique even globally. This region is characterized by extensive boreal forests and vast areas of tundra, intermingled by various freshwater bodies. Climate change and loss of natural habitats due to intensive land-use and unsustainable use of natural resources are the most serious threats to biodiversity in this region, also called the Barents Region.

Refugia, the sites preserving conditions reminiscent of suitable climates, are projected to be crucial for species in a changing climate, particularly at high latitudes. However, the knowledge of current locations of high-latitude refugia and particularly their ability to retain suitable conditions under future climatic changes is limited. Occurrences of refugia have previously been mainly assessed and modelled based solely on climatic features, with insufficient attention being paid to potentially important landscape-scale factors. Here, climate-only models and full' models incorporating topo-edaphic landscape-scale variables (radiation, soil moisture and calcareousness) were developed and compared for 111 arctic-alpine plant species in Northern Fennoscandia. This was done for both current and future climates to determine cells with resilient climatic suitability harbouring refugia. Our results show that topographic and edaphic landscape-scale predictors both significantly improve models of arctic-alpine species distributions and alter projections of refugia occurrence. The predictions of species-climate models ignore landscape-scale ecological processes and may thus provide inaccurate estimates of extinction risk and forecasts of refugia where species can persist under a changing climate.

Determining the drivers, patterns and hotspots of biodiversity can be of critical importance in supporting regional conservation planning. However, as biodiversity hotspots can be described with several different metrics, it is important to investigate their congruence as well as the spatial overlap of hotspots with protected areas. Here, by using extensive data on climate, topography, soil characteristics and vascular plants combined with boosted regression tree models, we determine the patterns and key drivers of plant diversity hotspots along broad environmental gradients in northernmost Europe spanning from taiga landscapes to treeless tundra. We assess plant diversity with four metrics – species richness, range-rarity richness, threatened species richness, and local contribution to beta diversity – and examine their congruence with each other as well as with contemporary conservation areas. We found that climate plays an important role in governing species diversity, though topo-edaphic are highlighted alongside climatic predictors in determining the diversity patterns of many threatened, near-threatened, and range-restricted species. Importantly, the different diversity metrics have contrasting drivers and, overall, their hotspots have low congruence. Furthermore, existing protected areas appear to offer limited coverage for hotspots of vascular plant diversity. Modelling the various facets of diversity and their drivers, such as the topo-edaphic setting, may provide useful information to help conserve diversity in a changing climate. Projected patterns of different aspects of diversity and their congruency can provide insights into the processes underlying biodiversity and be employed to assess the representativeness of protected area networks.

Aim Nitrogen deposition is a major global driver of change in plant communities, but its impacts on higher trophic levels are insufficiently understood. Here, we introduce and test a novel conceptual trait-based model describing how the effects of soil eutrophication cascade to higher trophic levels across differential plant–herbivore interactions. Location Northern Europe. Methods We synthesize previous literature on the effects of nitrogen on plants and herbivorous insects as well as relevant multispecies patterns of insect communities concerning species dietary breadth, body size, dispersal propensity and voltinism in order to derive the model. We empirically evaluate the proposed, hitherto untested, four main model pathways using statistical modelling and data on 1064 northern European butterfly and moth species, their life-history traits, phylogeny and population trends. Results We show that across all species: (1) larval dietary breadth and host plant foliar nitrogen content are positively and equally strongly related to insect body size, and that (2) multivoltinism, host plant preferences for soil nitrogen, body size and larval dietary breadth are positively related to population trends of butterflies and moths as predicted by the model. Positive relationships between plant foliar nitrogen content and body size as well as multivoltinism and population trends are the first multispecies demonstrations for these patterns. Main conclusions Soil nitrogen enrichment amplifies the diverging trends of herbivorous insects feeding on nitrophilous versus nitrophobous plants through differential plant–herbivore interactions, causing predictable changes in community composition at higher trophic levels. A positive foliar nitrogen–insect body size relationship, now empirically supported, is the integrating link within this cascade. As nitrogen deposition is a global driver, our model suggests that a major future trend may be an increased dominance of insects that are large, dispersive, multivoltine, dietary generalists or specialized on nitrophilous plant species at the expense of species preferring oligotrophic environments.

Assisted migration (AM) has been suggested as a management strategy for aiding species in reaching newly suitable locations as climate changes. Species distribution models (SDMs) can provide important insights for decisions on whether to assist a species in its migration; however, their application includes uncertainties. In this study, we use consensus SDMs to model the future suitable areas for 13 vascular plant species with poor dispersal capacity. Based on the outputs of SDMs under different climate change scenarios and future times, we quantify the predicted changes in suitable area by calculating metrics that describe the need and potential for migration. We find that, by the end of the 21st century, one of the species would benefit from AM under mild climate change, seven under moderate change, and for 12 out of 13 species studied AM appears to be a relevant conservation method under strong climate change. We also test the effect of different modeling attributes on the metrics and find little variation between SDMs constructed using different combinations of modeling methods and variable sets. However, the choice of climate variables had a larger influence on the level of the metrics than did the modeling method. We therefore suggest that the choice of climate variables should receive ample attention when measuring climate change threat using SDMs and that experiments aiming to uncover critical environmental factors for individual species should be extensively conducted. This study illustrates that dispersal assistance may be needed for many species under a wide range of possible future climates.

European aspen (Populus tremula L.) is a keystone species in boreal forests that are dominated by coniferous tree species. Both living and dead aspen trees contribute significantly to the species diversity of forest landscapes. Thus, spatial and temporal continuity of aspen is a prerequisite for the long-term persistence of viable populations of numerous aspen-associated species. In this review, we collate existing knowledge on the ecological role of European aspen, assess the knowledge needs for aspen occurrence patterns and dynamics in boreal forests and discuss the potential of different remote sensing techniques in mapping aspen at various spatiotemporal scales. The role of aspen as a key ecological feature has received significant attention, and studies have recognised the negative effects of modern forest management methods and heavy browsing on aspen occurrence and regeneration. However, the spatial knowledge of occurrence, abundance and temporal dynamics of aspen is scarce and incomprehensive. The remote sensing studies reviewed here highlight particularly the potential of three-dimensional data derived from airborne laser scanning or photogrammetric point clouds and airborne imaging spectroscopy in mapping European aspen, quaking aspen (Populus tremuloides Michx.) and other Populus species. In addition to tree species discrimination, these methods can provide information on biophysical, biochemical properties and even genetic diversity of aspen trees. Major obstacles in aspen detection using remote sensing are the low proportion and scattered occurrence of European aspen in boreal forests and the overlap of spectral and/or structural properties of European aspen and quaking aspen with some other tree species. Furthermore, the suitability of remote sensing data for aspen mapping and monitoring depends on the geographical coverage of data, the availability of multitemporal data and the costs of data acquisition. Our review highlights that integration of ecological knowledge with spatiotemporal information acquired by remote sensing is key to understanding the current and future distribution patterns of aspen-related biodiversity.

International audience; Predictive models are central to many scientific disciplines and vital for informing management in a rapidly changing world. However, limited understanding of the accuracy and precision of models transferred to novel conditions (their 'trans-ferability') undermines confidence in their predictions. Here, 50 experts identified priority knowledge gaps which, if filled, will most improve model transfers. These are summarized into six technical and six fundamental challenges, which underlie the combined need to intensify research on the determinants of ecological predictability, including species traits and data quality, and develop best practices for transferring models. Of high importance is the identification of a widely applicable set of transferability metrics, with appropriate tools to quantify the sources and impacts of prediction uncertainty under novel conditions. Predicting the Unknown Predictions facilitate the formulation of quantitative, testable hypotheses that can be refined and validated empirically [1]. Predictive models have thus become ubiquitous in numerous scientific disciplines, including ecology [2], where they provide means for mapping species distributions, explaining population trends, or quantifying the risks of biological invasions and disease outbreaks (e.g., [3,4]). The practical value of predictive models in supporting policy and decision making has therefore grown rapidly (Box 1) [5]. With that has come an increasing desire to predict (see Glossary) the state of ecological features (e.g., species, habitats) and our likely impacts upon them [5], prompting a shift from explanatory models to anticipatory predictions [2]. However, in many situations, severe data deficiencies preclude the development of specific models, and the collection of new data can be prohibitively costly or simply impossible [6]. It is in this context that interest in transferable models (i.e., those that can be legitimately projected beyond the spatial and temporal bounds of their underlying data [7]) has grown. Transferred models must balance the tradeoff between estimation and prediction bias and variance (homogenization versus nontransferability, sensu [8]). Ultimately, models that can Highlights Models transferred to novel conditions could provide predictions in data-poor scenarios, contributing to more informed management decisions.

Climate change is a major threat to biodiversity, causing species to move to new climatically suitable areas, and thus increasing the extinction probability of species inhabiting fragmented landscapes. This highlights the need for climate-wise conservation strategies. With such strategies, a well-connected network of protected areas (PAs) is one of the most important means to support species survival. An extensive and representative PA network can enhance the resilience of regional populations of species, resulting in slower species loss in landscapes with a significant proportion of area of habitat being protected. This paper presents analyses of both the observed (1974–2010) and the predicted changes (by 2051–2080) in boreal bird populations in Finland. Firstly, the results show some general patterns of climate change on bird species: (1) species are shifting their ranges towards north, (2) range sizes of many species are declining, and (3) these changes are different in northern and southern species and in species occupying different habitats. Secondly, the paper looks more into the role of protected area (PA) network in securing birds in a changing climate and concludes that at least in Finland, open habitats, such as open mires and mountain heaths, change more rapidly in their species composition in protected areas than for example old-growth forests. However, generally, species decline less within than outside PAs showing that protected areas alleviate climate change effects on bird species of conservation concern. This finding, further supported by results from elsewhere in Europe, provides evidence for the resilience of PA networks in preserving species under climate change. Representative PA network that includes high cover for key habitats is hence needed in all latitudinal zones. The projected efficiency of the PA network in maintaining biodiversity was partly dependent on the strength of climate change varying with respect to future scenarios. This suggests that a flexibly adaptive climate-wise conservation planning is required to be better prepared for preserving biodiversity in the face of uncertain climate change. Thirdly, the paper discusses several aspects of climate change studies and avian biodiversity that have been hitherto understudied especially in the northern biomes. The paper suggests that future studies should concentrate on (1) abundance-based models and prioritisations, (2) species’ adaptive capacity (ability to avoid the impacts of climate change through dispersal and/or evolutionary change) and sensitivity (limited potential to persist in situ under changing climate) to climate change, (3) the role of the landscape matrix around the PAs and (4) the effects of the biogeophysical features of the PAs themselves. In conclusion, we envision that improved assessments regarding the ability of PA networks to maintain biodiversity in northern biomes are needed to enhance our ability to perform climate-wise conservation planning.

Model transferability is an emerging and important branch of predictive science that has grown primarily from a need to produce ecological forecasts in the face of widespread data deficiency and escalating environmental novelty. In our recent article in Trends in Ecology and Evolution, we outlined some of the major roadblocks that currently undermine the practice of model transfers in ecology. The response of Radchuk et al. to our work stresses the value of considering ‘first principles’ in projections of ecosystem change and offers insights into outstanding challenges specific to mechanistic (synonym: process-based) models.

Aim Biotic interactions have a central role in defining species assemblages, realized both through negative and positive impacts. However, forecasts of how these interactions affect biodiversity across landscapes are challenging (and lacking) because the outcome of interactions depends not only on the identity of the interacting species but also on local environmental conditions. Thus, we study how biotic interactions manifest across a landscape. Location High-latitude northern Finland and Norway (69° N, 21° E). Methods We modelled the influence of a dominant shrub, Empetrum hermaphroditum, on spatial patterns of species richness of vascular plants, bryophytes and lichens (using cover of the shrub as a proxy for the frequency and intensity of interaction with other species) across a topographically variable landscape. Results The relationship between the cover of the dominant shrub and species richness differed between guilds, being strongest for vascular plant richness, and varied considerably along environmental gradients. The dominant shrub showed stronger negative impact on vascular plant richness under less abiotically extreme conditions. For lichens, the relationship was the opposite: under mild conditions, species richness increased with the cover of the shrub. Incorporating specific leaf area (SLA) data into analyses revealed that the dominant shrub affected species with high SLA (typically competitive species of resource-rich environments) most negatively. Main conclusions Our findings show how the impact of a dominant species varies across a landscape, with distinctly different effects on competitive and stress-tolerant vascular plants, and on bryophytes and lichens. Spatial predictions of biodiversity trends under global environmental change could, therefore, critically benefit from accounting for context-dependent impacts of biotic interactions.

Success rates for conservation translocations of species are low and there is a need for increased understanding of how this activity is best applied. Here, using static species distribution models and a spatially-explicit dynamic simulation model, RangeShifter, we examine the impacts of habitat cover in recipient landscapes, allocation of individuals into multiple sites and species trait characteristics on the long-term fate of hypothetical translocations of a grassland specialist butterfly, Maniola jurtina, in Finland. While persistence of populations introduced to climatically suitable locations northwards of the current range can be increased by selecting sites with increasing habitat cover and by allocation of individuals to multiple release sites, local population growth rate is shown to be the key parameter in determining likely translocation success. We conclude that the long-term persistence of translocated habitat specialist butterflies, particularly with low growth rates, appears to be uncertain in modern-day fragmented grassland networks and that translocation activities should prioritize management that improves local growth rate.

Biotic interactions may strongly affect the distribution of individual species and the resulting patterns of species richness. However, the impacts can vary depending on the species or taxa examined, suggesting that the influences of interactions on species distributions and diversity are not always straightforward and can be taxon-contingent. The aim of this study was therefore to examine how the importance of biotic interactions varies within a community. We incorporated three biotic predictors (cover of the dominant vascular species) into two correlative species richness modelling frameworks to predict spatial variation in the number of vascular plants, bryophytes and lichens in arctic–alpine Fennoscandia, in N Europe. In addition, predictions based on single-species distribution models were used to determine the nature of the impact (negative vs. positive outcome) of the three dominant species on individual vascular plant, bryophyte and lichen species. Our results suggest that biotic variables can be as important as abiotic variables, but their relative contributions in explaining the richness of sub-dominant species vary among dominant species, species group and the modelling framework implemented. Similarly, the impacts of biotic interactions on individual species varied among the three species groups and dominant species, with the observed patterns partly reflecting species’ biogeographic range. Our study provides additional support for the importance of biotic interactions in modifying arctic–alpine biodiversity patterns and highlights that the impacts of interactions are not constant across taxa or biotic drivers. The influence of biotic interactions, including the taxon contingency and range-based impacts, should therefore be accounted for when developing biodiversity forecasts.

Translocations have been advocated as a conservation tool helping species adapt to climate and land-use change, but well-documented examples of invertebrates’ translocations are rare. The paper describes a successful translocation of the threatened Clouded Apollo butterfly ( Parnassius mnemosyne ) in Finland, compares this to a specific failed translocation, and presents conclusions for conservation planning as to factors contributing to the success. Two apparent key characteristics of the successful translocation were greater abundance of larval resources and less open landscape. The successful site was surrounded by forest, which strongly restricted emigration, crucially supporting the survival of the small initial population. Based on 20 mated females’ translocation in 2000, the local population increased slowly, reaching 600 butterflies in 2011. A large translocation patch together with host-plant abundance enabled successful establishment of the local population. Availability of other suitable grassland patches sufficiently nearby was an additional key characteristic, facilitating the Clouded Apollo’s expansion. However, the expansion rate was low; it took seven years for the butterflies to colonise the five nearest patches, only 10–200 m from the translocation patch. By 2013, they had colonised all suitable semi-natural grassland patches within 2 km from the translocation site and established a seemingly viable metapopulation with 11 subpopulations. The results point to the significance of local habitat area and landscape quality, along with conditions restricting emigration, in determination of suitable translocation sites.

Projected climatic warming calls for increased attention to the identification of suitable refugia for the preservation of biota and ecosystems in changing high-latitude environments. One such way is the development of models for drivers of refugia. Here, we investigate the distribution and species richness of Arctic-alpine vascular plant species' refugia. The study is carried out in an environmentally variable area in N Europe, encompassing the northern boreal to the Arctic-alpine zone. We defined refugia as isolated 1 km x 1 km grid cells with multiple Arctic-alpine plant species occurrences outside their main distribution area and assessed the main environmental factors underlying their distribution and richness using cross-validated boosted regression tree modelling. In the modelling, we examined the effects of climatic, topographic, and geologic factors, and the connectivity of sites with refugia incrementally, i.e. first modelling climatic impact alone, then with separate additions of topographic, geologic and connectivity variables, concluding with a model including all predictor variables. The inclusion of slope and connectivity significantly improved model performance. Although climate has a central role in controlling the occurrence of refugia, topography provides important clues for recognizing heterogeneous locations that harbour refugia with suitable local thermal and moisture conditions. Results suggest considering refugia as, on the one hand, isolated pockets of suitable habitat, but on the other hand as potentially interconnected habitat networks. In general, our study demonstrates that the spatial patterns of refugia can be successfully modelled, but emphasizes a need for high-quality data sampled at resolutions reflecting significant environmental gradients.

Climatically driven Earth surface processes (ESPs) govern landscape and ecosystem dynamics in high-latitude regions. However, climate change is expected to alter ESP activity at yet uncertain rate and amplitude. We examined the sensitivity of key ESPs (cryoturbation, solifluction, nivation, and palsa mires) to changing climate by modeling their distribution in regard to climate, local topography, and soil variables in northern Fennoscandia. The distributions of ESPs were then forecasted under two future time periods, 2040–2069 and 2070–2099, using ensemble modeling and three emission scenarios. Increase of 2°C in current temperature conditions caused an almost complete loss of ESPs, highlighting the extreme climatic sensitivity of high-latitude geomorphic processes. Forecasts based on three scenarios suggest a disappearance of suitable climate for studied ESPs by the end of this century. This could initiate multiple opposing feedback between land surface and atmosphere through changes in albedo, heat fluxes, and biogeochemical cycles.

Summary Rapid environmental changes are threatening biodiversity and exposing species to novel ecological and evolutionary pressures. The scientific community increasingly recognizes the need for dynamic models integrating sufficient complexity both to improve our understanding of species' responses to environmental changes and to inform effective management strategies. Using three illustrative examples, we introduce a novel modelling platform, RangeShifter, which integrates complex population dynamics and dispersal behaviour, includes plastic and evolutionary processes and simulates scenarios on spatially explicit landscapes. The software provides functionality for a wide variety of modelling applications ranging from applied questions, where it can be parameterized for real landscapes and species to compare alternative potential management interventions, to purely theoretical studies of species' eco-evolutionary dynamics and responses to different environmental pressures. RangeShifter provides an important tool for facilitating the advancement of ecological theory on species' spatial dynamics in response to environmental changes, and linking it directly to application in biodiversity conservation.

Species inhabiting high-latitude environments are anticipated to be exceptionally vulnerable to climate change because of the greater temperature increases projected for these regions. Earlier studies based on bioclimatic envelope models and bird atlas data from 1974 to 1989 have suggested that northern-boreal bird species may face considerable range contractions in Northern Europe by 2080. Using new bird atlas data from Finland compiled in 2006–2010, we show that the ranges of these northern-boreal bird species have already contracted by 27%, on average, compared with 1974–89. The majority (21) of the studied 27 northern-boreal bird species showed significant contractions of ranges and/or northwards shifts. Thus the range changes of northern species are in the same direction as the predictions of species-climate change models.

Remote sensing (RS) data may play an important role in the development of cost-effective means for modelling, mapping, planning and conserving biodiversity. Specifically, at the landscape scale, spatial models for the occurrences of species of conservation concern may be improved by the inclusion of RS-based predictors, to help managers to better meet different conservation challenges. In this study, we examine whether predicted distributions of 28 red-listed plant species in north-eastern Finland at the resolution of 25 ha are improved when advanced RS-variables are included as unclassified continuous predictor variables, in addition to more commonly used climate and topography variables. Using generalized additive models (GAMs), we studied whether the spatial predictions of the distribution of red-listed plant species in boreal landscapes are improved by incorporating advanced RS (normalized difference vegetation index, normalized difference soil index and Tasseled Cap transformations) information into species-environment models. Models were fitted using three different sets of explanatory variables: (1) climate-topography only; (2) remote sensing only; and (3) combined climate-topography and remote sensing variables, and evaluated by four-fold cross-validation with the area under the curve (AUC) statistics. The inclusion of RS variables improved both the explanatory power (on average 8.1 % improvement) and cross-validation performance (2.5 %) of the models. Hybrid models produced ecologically more reliable distribution maps than models using only climate-topography variables, especially for mire and shore species. In conclusion, Landsat ETM+ data integrated with climate and topographical information has the potential to improve biodiversity and rarity assessments in northern landscapes, especially in predictive studies covering extensive and remote areas.

Aim To quantify whether species distribution models (SDMs) can reliably forecast species distributions under observed climate change. In particular, to test whether the predictive ability of SDMs depends on species traits or the inclusion of land cover and soil type, and whether distributional changes at expanding range margins can be predicted accurately. Location Finland Methods Using 10-km resolution butterfly atlas data from two periods, 1992–99 (t1) and 2002–09 (t2), with a significant between-period temperature increase, we modelled the effects of climatic warming on butterfly distributions with boosted regression trees (BRTs) and generalized additive models (GAMs). We evaluated model performance by using the split-sample approach with data from t1 (‘nonindependent validation’), and then compared model projections based on data from t1 with species’ observed distributions in t2 (‘independent validation’). We compared climate-only SDMs to SDMs including land cover, soil type, or both. Finally, we related model performance to species traits and compared observed and predicted distributional shifts at northern range margins.

Amphibian declines have been reported worldwide during the last decades. In this study, we focused on the endangered great crested newt (Triturus cristatus), which has suffered from intensive forestry and past mire ditching in the northern verge of its distribution. We collected data from 46 breeding ponds in eastern Finland during 2005–2011 using dip-netting. We modeled breeding success with cost-effective methods by using site-level forest data. The results highlight the importance of herb-rich forests in the vicinity of breeding ponds. Based on the results possible new breeding sites can now be located based on the presence and size of types of nutrient-rich forest, proportion of deciduous trees, young stands, shading and unfavorable habitats in the vicinity of ponds. Also practical conservation measures, like creating new ponds within a certain area, can now be allocated more accurately using the same variables.

Agricultural intensification is a major driver of biodiversity decline throughout Europe. Agri-environment schemes governed by EU regulation are a significant tool in combating this decline but despite high spending, experiences of their effectiveness have been mixed. Their effectiveness might be improved by targeting them to locations with high biodiversity value, and particularly by spatial coordination to enhance habitat connectivity and the associated ecological processes, such as dispersal. We show, with an example of semi-natural grassland conservation in South-Western Finland, how spatial conservation planning tools, here the Zonation software, could help in assessing the habitat connectivity and allocating management actions. We assign highest priority to sites that have been classified as nationally important and that have been under management, and let connectivity influence the Zonation prioritizations. According to Zonation outputs, 25–30% of highest-ranking grasslands in our study area are without management contracts, indicating weak connectivity of managed sites, whereas Natura 2000 areas are spatially better located. A 50% expansion of the current network would be adequate to bring its value close to that of a network created from scratch, but as the contracts are temporary, reallocation of the contracts from the least valuable, funded sites to more valuable, unfunded ones would be even more effective. Current policy instruments supporting farmland biodiversity are strongly constrained by EU regulation, and appear to be too inflexible to take the spatial differences in conservation values into account. Better communication and incentives to encourage farmer participation to these voluntary programs are needed.

Species ranges are expected to move polewards following the changing climate, which poses novel challenges to the protected area network, particularly at northern latitudes. Here we study how well protected areas are likely to sustain populations of birds of conservation concern under a changing climate in northern Europe, in Finland. We fitted bioclimatic envelope models generated for 100 bird species to climate scenario data for the years 2051–2080 and three alternative emission scenarios in a 10-km grid system to predict changes in the species probability of occurrence. We related the projected changes in the climatic suitability to the amount of protected preferred habitat for the study species in the 10-km grid cells, and based on the cover of four main CORINE Land Cover classes in each conservation area in Finland. The probability of occurrence of all species (except marshland birds) decreased according to all scenarios, the decline being greatest in southern and smallest in northern boreal zones. This decline was slightly greater in unprotected than in protected areas for species of forests, mires and mountain habitats. The climatically suitable areas for the species were predicted to shift northwards, but the potential gain of southern species of conservation concern appears not to compensate for the loss of northern species. Thus, a representative protected area network is needed in all boreal zones. Overall, our results show that species-specific habitat preferences and habitat availability should be taken into account when assessing the efficiency of a protected area network in a changing climate.

Measures of geodiversity may provide a potentially useful surrogate for biodiversity patterns in insufficiently surveyed areas. However, their reliability in modelling the spatial variation in species richness is inadequately understood. We investigated whether the explanatory and predictive power of species richness models can be improved by considering explicit measures of geodiversity (variability of earth surface materials, forms and processes) in addition to climate and topography variables. Vascular plant species richness was modelled in two study areas in Northern Europe, Finland at the resolution of 500 or 1000 m, and as a function of three geodiversity (geological, geomorphological and hydrological diversity) variables, and six climate and topography variables. Variation partitioning was used to identify the independent and shared contributions of the geodiversity, climate and topography variable groups in explaining the spatial patterns of species richness. Generalized additive models were used to explore the ability of the different explanatory variables in predicting plant species richness within and between the study areas. In both the study areas, the inclusion of measures of geodiversity improved the explanatory power, predictive ability and robustness of the plant species richness models. In conclusion, the explicit measures of geodiversity appear to be promising surrogates of biodiversity, which reflect important abiotic resource factors, and may thus provide an equally, or even more reliable basis for transferring biodiversity models to new areas than models based on climate and topography variables.