Recent range shifts of moths, butterflies, and birds are driven by the breadth of their climatic niche.

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.

Role of data uncertainty when identifying important areas for biodiversity and carbon in boreal forests.

Forest conservation plays a central role in meeting national and international biodiversity and climate targets. Biodiversity and carbon values within forests are often estimated with models, introducing uncertainty to decision making on which forest stands to protect. Here, we explore how uncertainties in forest variable estimates affect modelled biodiversity and carbon patterns, and how this in turn introduces variability in the selection of new protected areas. We find that both biodiversity and carbon patterns were sensitive to alterations in forest attributes. Uncertainty in features that were rare and/or had dissimilar distributions with other features introduced most variation to conservation plans. The most critical data uncertainty also depended on what fraction of the landscape was being protected. Forests of highest conservation value were more robust to data uncertainties than forests of lesser conservation value. Identifying critical sources of model uncertainty helps to effectively reduce errors in conservation decisions.

Effect of forest management choices on carbon sequestration and biodiversity at national scale.

Forest management methods and harvest intensities influence wood production, carbon sequestration and biodiversity. We devised different management scenarios by means of stakeholder analysis and incorporated them in the forest growth simulator PREBAS. To analyse impacts of harvest intensity, we used constraints on total harvest: business as usual, low harvest, intensive harvest and no harvest. We carried out simulations on a wall-to-wall grid in Finland until 2050. Our objectives were to (1) test how the management scenarios differed in their projections, (2) analyse the potential wood production, carbon sequestration and biodiversity under the different harvest levels, and (3) compare different options of allocating the scenarios and protected areas. Harvest level was key to carbon stocks and fluxes regardless of management actions and moderate changes in proportion of strictly protected forest. In contrast, biodiversity was more dependent on other management variables than harvesting levels, and relatively independent of carbon stocks and fluxes.

Modelling the regional potential for reaching carbon neutrality in Finland: Sustainable forestry, energy use and biodiversity protection.

The EU aims at reaching carbon neutrality by 2050 and Finland by 2035. We integrated results of three spatially distributed model systems (FRES, PREBAS, Zonation) to evaluate the potential to reach this goal at both national and regional scale in Finland, by simultaneously considering protection targets of the EU biodiversity (BD) strategy. Modelling of both anthropogenic emissions and forestry measures were carried out, and forested areas important for BD protection were identified based on spatial prioritization. We used scenarios until 2050 based on mitigation measures of the national climate and energy strategy, forestry policies and predicted climate change, and evaluated how implementation of these scenarios would affect greenhouse gas fluxes, carbon storages, and the possibility to reach the carbon neutrality target. Potential new forested areas for BD protection according to the EU 10% protection target provided a significant carbon storage (426-452 TgC) and sequestration potential (- 12 to - 17.5 TgCO2eq a-1) by 2050, indicating complementarity of emission mitigation and conservation measures. The results of the study can be utilized for integrating climate and BD policies, accounting of ecosystem services for climate regulation, and delimitation of areas for conservation.

Bioclimate change across the protected area network of Finland.

Protected areas (PAs) are crucial in conserving biodiversity under climate change. In boreal regions, trends of biologically relevant climate variables (i.e., bioclimate) in PAs have remained unquantified. We investigated the changes and variability of 11 key bioclimatic variables across Finland during the period 1961-2020 based on gridded climatology. Our results suggest significant changes in annual mean and growing season temperatures over the entire study area, whereas, e.g., annual precipitation sum and April-September water balance have increased especially in the central and northern parts of Finland. We found substantial variation in bioclimatic changes over the 631 studied PAs; in the northern boreal zone (NB) the number of snow-covered days has decreased on average by 5.9 days between 1961-1990 and 1991-2020, while in the southern boreal zone (SB) the corresponding decrease has been 16.1 days. The number of frost days in spring with absent snow cover has decreased in the NB (on average -0.9 days) while increasing in the SB (0.5 days), reflecting the changing exposure of biota to frost. The observed increases in accumulation of heat in the SB and more frequent rain-on-snow events in the NB can affect drought tolerance and winter survival of species, respectively. Principal component analysis suggested that the main dimensions of bioclimate change in PAs vary across vegetation zones; for example, in the SB the changes are related to annual and growing season temperatures, whereas in the middle boreal zone the changes are linked to altered moisture and snow conditions. Our results highlight the substantial spatial variation in bioclimatic trends and climate vulnerability across the PAs and vegetation zones. These findings provide a basis for the understanding of the multifaceted changes the boreal PA network is facing and help to develop and direct conservation and management.

Developing fine-grained nationwide predictions of valuable forests using biodiversity indicator bird species.

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 × 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.

Decreasing snow cover alters functional composition and diversity of Arctic tundra.

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.

Fine-grained climate velocities reveal vulnerability of protected areas to climate change.

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.

Outstanding Challenges in the Transferability of Ecological Models.

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 'transferability') 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.

Weather explains high annual variation in butterfly dispersal.

Weather conditions fundamentally affect the activity of short-lived insects. Annual variation in weather is therefore likely to be an important determinant of their between-year variation in dispersal, but conclusive empirical studies are lacking. We studied whether the annual variation of dispersal can be explained by the flight season's weather conditions in a Clouded Apollo (Parnassius mnemosyne) metapopulation. This metapopulation was monitored using the mark-release-recapture method for 12 years. Dispersal was quantified for each monitoring year using three complementary measures: emigration rate (fraction of individuals moving between habitat patches), average residence time in the natal patch, and average distance moved. There was much variation both in dispersal and average weather conditions among the years. Weather variables significantly affected the three measures of dispersal and together with adjusting variables explained 79-91% of the variation observed in dispersal. Different weather variables became selected in the models explaining variation in three dispersal measures apparently because of the notable intercorrelations. In general, dispersal rate increased with increasing temperature, solar radiation, proportion of especially warm days, and butterfly density, and decreased with increasing cloudiness, rainfall, and wind speed. These results help to understand and model annually varying dispersal dynamics of species affected by global warming.

Protected areas alleviate climate change effects on northern bird species of conservation concern.

Global climate change is a major threat to biodiversity, posing increasing pressures on species to adapt in situ or shift their ranges. A protected area network is one of the main instruments to alleviate the negative impacts of climate change. Importantly, protected area networks might be expected to enhance the resilience of regional populations of species of conservation concern, resulting in slower species loss in landscapes with a significant amount of protected habitat compared to unprotected landscapes. Based on national bird atlases compiled in 1974-1989 and 2006-2010, this study examines the recent range shifts in 90 forest, mire, marshland, and Arctic mountain heath bird species of conservation concern in Finland, as well as the changes in their species richness in protected versus unprotected areas. The trends emerging from the atlas data comparisons were also related to the earlier study dealing with predictions of distributional changes for these species for the time slice of 2051-2080, developed using bioclimatic envelope models (BEMs). Our results suggest that the observed changes in bird distributions are in the same direction as the BEM-based predictions, resulting in a decrease in species richness of mire and Arctic mountain heath species and an increase in marshland species. The patterns of changes in species richness between the two time slices are in general parallel in protected and unprotected areas. However, importantly, protected areas maintained a higher level of species richness than unprotected areas. This finding provides support for the significance and resilience provision of protected area networks in preserving species of conservation concern under climate change.

Impacts of land cover data selection and trait parameterisation on dynamic modelling of species' range expansion.

Dynamic models for range expansion provide a promising tool for assessing species' capacity to respond to climate change by shifting their ranges to new areas. However, these models include a number of uncertainties which may affect how successfully they can be applied to climate change oriented conservation planning. We used RangeShifter, a novel dynamic and individual-based modelling platform, to study two potential sources of such uncertainties: the selection of land cover data and the parameterization of key life-history traits. As an example, we modelled the range expansion dynamics of two butterfly species, one habitat specialist (Maniola jurtina) and one generalist (Issoria lathonia). Our results show that projections of total population size, number of occupied grid cells and the mean maximal latitudinal range shift were all clearly dependent on the choice made between using CORINE land cover data vs. using more detailed grassland data from three alternative national databases. Range expansion was also sensitive to the parameterization of the four considered life-history traits (magnitude and probability of long-distance dispersal events, population growth rate and carrying capacity), with carrying capacity and magnitude of long-distance dispersal showing the strongest effect. Our results highlight the sensitivity of dynamic species population models to the selection of existing land cover data and to uncertainty in the model parameters and indicate that these need to be carefully evaluated before the models are applied to conservation planning.

Climate change, northern birds of conservation concern and matching the hotspots of habitat suitability with the reserve network.

National reserve networks are one of the most important means of species conservation, but their efficiency may be diminished due to the projected climatic changes. Using bioclimatic envelope models and spatial data on habitats and conservation areas, we studied how efficient the reserve network will be in preserving 100 forest, mire, marshland, and alpine bird species of conservation concern in Finland in 2051-2080 under three different climate scenarios. The occurrences of the studied bird species were related to the amount of habitat preferred by each species in the different boreal zones. We employed a novel integrated habitat suitability index that takes into account both the species' probability of occurrence from the bioclimatic models and the availability of suitable habitat. Using this suitability index, the distribution of the topmost 5% suitability squares ("hotspots") in the four bird species groups in the period 1971-2000 and under the three scenarios were compared with the location of reserves with the highest amounts of the four habitats to study the efficiency of the network. In species of mires, marshlands, and Arctic mountains, a high proportion of protected habitat was included in the 5% hotspots in the scenarios in 2051-2080, showing that protected areas cover a high proportion of occurrences of bird species. In contrast, in forests in the southern and middle boreal zones, only a small proportion of the protected habitat was included in the 5% hotspots, indicating that the efficiency of the protected area network will be insufficient for forest birds in the future. In the northern boreal zone, the efficiency of the reserve network in forests was highly dependent on the strength of climate change varying between the scenarios. Overall, there is no single solution to preserving biodiversity in a changing climate, but several future pathways should be considered.

The role of biotic interactions in shaping distributions and realised assemblages of species: implications for species distribution modelling.

Predicting which species will occur together in the future, and where, remains one of the greatest challenges in ecology, and requires a sound understanding of how the abiotic and biotic environments interact with dispersal processes and history across scales. Biotic interactions and their dynamics influence species' relationships to climate, and this also has important implications for predicting future distributions of species. It is already well accepted that biotic interactions shape species' spatial distributions at local spatial extents, but the role of these interactions beyond local extents (e.g. 10 km(2) to global extents) are usually dismissed as unimportant. In this review we consolidate evidence for how biotic interactions shape species distributions beyond local extents and review methods for integrating biotic interactions into species distribution modelling tools. Drawing upon evidence from contemporary and palaeoecological studies of individual species ranges, functional groups, and species richness patterns, we show that biotic interactions have clearly left their mark on species distributions and realised assemblages of species across all spatial extents. We demonstrate this with examples from within and across trophic groups. A range of species distribution modelling tools is available to quantify species environmental relationships and predict species occurrence, such as: (i) integrating pairwise dependencies, (ii) using integrative predictors, and (iii) hybridising species distribution models (SDMs) with dynamic models. These methods have typically only been applied to interacting pairs of species at a single time, require a priori ecological knowledge about which species interact, and due to data paucity must assume that biotic interactions are constant in space and time. To better inform the future development of these models across spatial scales, we call for accelerated collection of spatially and temporally explicit species data. Ideally, these data should be sampled to reflect variation in the underlying environment across large spatial extents, and at fine spatial resolution. Simplified ecosystems where there are relatively few interacting species and sometimes a wealth of existing ecosystem monitoring data (e.g. arctic, alpine or island habitats) offer settings where the development of modelling tools that account for biotic interactions may be less difficult than elsewhere.

Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels.

Intensification or abandonment of agricultural land use has led to a severe decline of semi-natural habitats across Europe. This can cause immediate loss of species but also time-delayed extinctions, known as the extinction debt. In a pan-European study of 147 fragmented grassland remnants, we found differences in the extinction debt of species from different trophic levels. Present-day species richness of long-lived vascular plant specialists was better explained by past than current landscape patterns, indicating an extinction debt. In contrast, short-lived butterfly specialists showed no evidence for an extinction debt at a time scale of c. 40 years. Our results indicate that management strategies maintaining the status quo of fragmented habitats are insufficient, as time-delayed extinctions and associated co-extinctions will lead to further biodiversity loss in the future.

Extinction debt: a challenge for biodiversity conservation.

Local extinction of species can occur with a substantial delay following habitat loss or degradation. Accumulating evidence suggests that such extinction debts pose a significant but often unrecognized challenge for biodiversity conservation across a wide range of taxa and ecosystems. Species with long generation times and populations near their extinction threshold are most likely to have an extinction debt. However, as long as a species that is predicted to become extinct still persists, there is time for conservation measures such as habitat restoration and landscape management. Standardized long-term monitoring, more high-quality empirical studies on different taxa and ecosystems and further development of analytical methods will help to better quantify extinction debt and protect biodiversity.

New insights into butterfly-environment relationships using partitioning methods.

Variation partitioning and hierarchical partitioning are novel statistical approaches that provide deeper understanding of the importance of different explanatory variables for biodiversity patterns than traditional regression methods. Using these methods, the variation in occupancy and abundance of the clouded apollo butterfly (Parnassius mnemosyne L.) was decomposed into independent and joint effects of larval and adult food resources, microclimate and habitat quantity. The independent effect of habitat quantity variables (habitat area and connectivity) captured the largest fraction of the variation in the clouded apollo patterns, but habitat connectivity had a major contribution only for occupancy data. The independent effects of resources and microclimate were higher on butterfly abundance than on occupancy. However, a considerable amount of variation in the butterfly patterns was accounted for by the joint effects of predictors and may thus be causally related to two or all three groups of variables. Abundance of the butterfly in the surroundings of the focal grid cell had a significant effect in all analyses, independently of the effects of other predictors. Our results encourage wider applications of partitioning methods in biodiversity studies.