Ecological barriers mediate spatiotemporal shifts of bird communities at a continental scale.

Species' range shifts and local extinctions caused by climate change lead to community composition changes. At large spatial scales, ecological barriers, such as biome boundaries, coastlines, and elevation, can influence a community's ability to shift in response to climate change. Yet, ecological barriers are rarely considered in climate change studies, potentially hindering predictions of biodiversity shifts. We used data from two consecutive European breeding bird atlases to calculate the geographic distance and direction between communities in the 1980s and their compositional best match in the 2010s and modeled their response to barriers. The ecological barriers affected both the distance and direction of bird community composition shifts, with coastlines and elevation having the strongest influence. Our results underscore the relevance of combining ecological barriers and community shift projections for identifying the forces hindering community adjustments under global change. Notably, due to (macro)ecological barriers, communities are not able to track their climatic niches, which may lead to drastic changes, and potential losses, in community compositions in the future.

Farmland practices are driving bird population decline across Europe.

Declines in European bird populations are reported for decades but the direct effect of major anthropogenic pressures on such declines remains unquantified. Causal relationships between pressures and bird population responses are difficult to identify as pressures interact at different spatial scales and responses vary among species. Here, we uncover direct relationships between population time-series of 170 common bird species, monitored at more than 20,000 sites in 28 European countries, over 37 y, and four widespread anthropogenic pressures: agricultural intensification, change in forest cover, urbanisation and temperature change over the last decades. We quantify the influence of each pressure on population time-series and its importance relative to other pressures, and we identify traits of most affected species. We find that agricultural intensification, in particular pesticides and fertiliser use, is the main pressure for most bird population declines, especially for invertebrate feeders. Responses to changes in forest cover, urbanisation and temperature are more species-specific. Specifically, forest cover is associated with a positive effect and growing urbanisation with a negative effect on population dynamics, while temperature change has an effect on the dynamics of a large number of bird populations, the magnitude and direction of which depend on species' thermal preferences. Our results not only confirm the pervasive and strong effects of anthropogenic pressures on common breeding birds, but quantify the relative strength of these effects stressing the urgent need for transformative changes in the way of inhabiting the world in European countries, if bird populations shall have a chance of recovering.

Temperature niche composition change inside and outside protected areas under climate warming.

Conservation of biodiversity relies heavily on protected areas but their role and effectiveness under a warming climate is still debated. We estimated the climate-driven changes in the temperature niche compositions of bird communities inside and outside protected areas in southern Canada. We hypothesized that communities inside protected areas include a higher proportion of cold-dwelling species than communities outside protected areas. We also hypothesized that communities shift to warm-dwelling species more slowly inside protected areas than outside. To study community changes, we used large-scale and long-term (1997-2019) data from the Breeding Bird Survey of Canada. To describe the temperature niche compositions of bird communities, we calculated the community temperature index (CTI) annually for each community inside and outside protected areas. Generally, warm-dwelling species dominated communities with high CTI values. We modeled temporal changes in CTI as a function of protection status with linear mixed-effect models. We also determined which species contributed most to the temporal changes in CTI with a jackknife approach. As anticipated, CTI was lower inside protected areas than outside. However, contrary to our expectation, CTI increased faster over time inside than outside protected areas and warm-dwelling species contributed most to CTI change inside protected areas. These results highlight the ubiquitous impacts of climate warming. Currently, protected areas can aid cold-dwelling species by providing habitat, but as the climate warms, the communities' temperature compositions inside protected areas quickly begin to resemble those outside protected areas, suggesting that protected areas delay the impacts of climate warming on cold-dwelling species.

Cambios en la composición del nicho térmico dentro y fuera de las áreas protegidas bajo el calentamiento climático Resumen La conservación de la biodiversidad depende mucho de las áreas protegidas, aunque todavía se debate su papel y efectividad bajo el calentamiento climático. Estimamos los cambios causados por el clima en la composición de los nichos térmicos de las comunidades de aves dentro y fuera de las áreas protegidas al sur de Canadá. Supusimos que las comunidades dentro de las áreas protegidas incluyen una proporción mayor de especies de zonas frías que las comunidades fuera de las áreas protegidas. También supusimos que las comunidades cambian a especies de zonas cálidas de forma más lenta dentro de las áreas protegidas que fuera de ellas. Usamos datos de gran escala y largo plazo (1997-2019) del Censo de Aves Reproductoras de Canadá para estudiar los cambios comunitarios. Calculamos el índice anual de temperatura comunitaria (ITC) para cada comunidad dentro y fuera de las áreas protegidas para describir las composiciones del nicho térmico de las comunidades de aves. En general, las especies de zonas cálidas dominaron las comunidades con valores altos del ITC. Simulamos los cambios temporales en el ITC como función del estado de protección mediante modelos lineales de efecto mixto. También determinamos cuáles especies contribuyen más a los cambios temporales en el ITC con un enfoque jackknife. Como lo anticipamos, el ITC fue menor dentro de las áreas protegidas que afuera. Sin embargo, contrario a nuestra hipótesis, el ITC incrementó más rápido con el tiempo dentro de las áreas protegidas y las especies de zonas cálidas contribuyeron más al cambio en el ITC también dentro de las áreas protegidas. Estos resultados resaltan el impacto universal del calentamiento climático. Actualmente, las áreas protegidas pueden auxiliar a las especies de zonas frías al proporcionarles hábitats, pero conforme la temperatura aumenta, las composiciones térmicas de las comunidades dentro de las áreas protegidas se asemejan rápidamente a aquellas fuera de las áreas protegidas, lo que sugiere que las áreas protegidas retrasan el impacto del calentamiento climático sobre las especies de zonas frías.

Shifts in timing and duration of breeding for 73 boreal bird species over four decades.

Breeding timed to match optimal resource abundance is vital for the successful reproduction of species, and breeding is therefore sensitive to environmental cues. As the timing of breeding shifts with a changing climate, this may not only affect the onset of breeding but also its termination, and thus the length of the breeding period. We use an extensive dataset of over 820K nesting records of 73 bird species across the boreal region in Finland to probe for changes in the beginning, end, and duration of the breeding period over four decades (1975 to 2017). We uncover a general advance of breeding with a strong phylogenetic signal but no systematic variation over space. Additionally, 31% of species contracted their breeding period in at least one bioclimatic zone, as the end of the breeding period advanced more than the beginning. We did not detect a statistical difference in phenological responses of species with combinations of different migratory strategy or number of broods. Nonetheless, we find systematic differences in species responses, as the contraction in the breeding period was found almost exclusively in resident and short-distance migrating species, which generally breed early in the season. Overall, changes in the timing and duration of reproduction may potentially lead to more broods co-occurring in the early breeding season-a critical time for species' reproductive success. Our findings highlight the importance of quantifying phenological change across species and over the entire season to reveal shifts in the community-level distribution of bird reproduction.

Decomposing the spatial and temporal effects of climate on bird populations in northern European mountains.

The relationships between species abundance or occurrence versus spatial variation in climate are commonly used in species distribution models to forecast future distributions. Under "space-for-time substitution", the effects of climate variation on species are assumed to be equivalent in both space and time. Two unresolved issues of space-for-time substitution are the time period for species' responses and also the relative contributions of rapid- versus slow reactions in shaping spatial and temporal responses to climate change. To test the assumption of equivalence, we used a new approach of climate decomposition to separate variation in temperature and precipitation in Fennoscandia into spatial, temporal, and spatiotemporal components over a 23-year period (1996-2018). We compiled information on land cover, topography, and six components of climate for 1756 fixed route surveys, and we modeled annual counts of 39 bird species breeding in the mountains of Fennoscandia. Local abundance of breeding birds was associated with the spatial components of climate as expected, but the temporal and spatiotemporal climatic variation from the current and previous breeding seasons were also important. The directions of the effects of the three climate components differed within and among species, suggesting that species can respond both rapidly and slowly to climate variation and that the responses represent different ecological processes. Thus, the assumption of equivalent species' response to spatial and temporal variation in climate was seldom met in our study system. Consequently, for the majority of our species, space-for-time substitution may only be applicable once the slow species' responses to a changing climate have occurred, whereas forecasts for the near future need to accommodate the temporal components of climate variation. However, appropriate forecast horizons for space-for-time substitution are rarely considered and may be difficult to reliably identify. Accurately predicting change is challenging because multiple ecological processes affect species distributions at different temporal scales.

Protected area characteristics that help waterbirds respond to climate warming.

Protected area networks help species respond to climate warming. However, the contribution of a site's environmental and conservation-relevant characteristics to these responses is not well understood. We investigated how composition of nonbreeding waterbird communities (97 species) in the European Union Natura 2000 (N2K) network (3018 sites) changed in response to increases in temperature over 25 years in 26 European countries. We measured community reshuffling based on abundance time series collected under the International Waterbird Census relative to N2K sites' conservation targets, funding, designation period, and management plan status. Waterbird community composition in sites explicitly designated to protect them and with management plans changed more quickly in response to climate warming than in other N2K sites. Temporal community changes were not affected by the designation period despite greater exposure to temperature increase inside late-designated N2K sites. Sites funded under the LIFE program had lower climate-driven community changes than sites that did not received LIFE funding. Our findings imply that efficient conservation policy that helps waterbird communities respond to climate warming is associated with sites specifically managed for waterbirds.

Las redes de áreas protegidas ayudan a las especies a responder al calentamiento climático. Sin embargo, se sabe muy poco sobre la contribución de las características ambientales y relevantes para la conservación de un sitio a estas respuestas. Investigamos cómo la composición de las comunidades no reproductivas de aves acuáticas (97 especies) en la red (3,018 sitios) Natura 2000 de la Unión Europea (N2K) cambió en respuesta a los incrementos de la temperatura durante más de 25 años en 26 países europeos. Medimos la reorganización comunitaria con base en series temporales de abundancia recolectadas durante el Censo Internacional de Aves Acuáticas en relación con los objetivos de conservación de los sitios N2K, el periodo de asignación de fondos y el estado del plan de manejo. La composición comunitaria de las aves acuáticas en los sitios con planes de manejo y designados explícitamente para su protección cambió más rápidamente en respuesta al calentamiento climático que en otros sitios N2K. Los cambios comunitarios temporales no se vieron afectados por el periodo de asignación a pesar de una mayor exposición al incremento de la temperatura dentro de los sitios N2K de asignación tardía. Los sitios financiados por el programa LIFE tuvieron menos cambios comunitarios causados por el clima que los sitios que no recibieron este financiamiento. Nuestros hallazgos sugieren que la política de conservación eficiente que ayuda a las comunidades de aves acuáticas a responder al calentamiento climático está asociada con sitios específicamente gestionados para las aves acuáticas.

Range shifts of overwintering birds depend on habitat type, snow conditions and habitat specialization.

Climatic warming is forcing species to shift their ranges poleward, which has been demonstrated for many taxa globally. Yet, the influence of habitat types on within- and among-species variations of distribution shifts has rarely been studied, especially during the non-breeding season. Here, we investigated habitat-specific shift distances of northern range margins and directions of the distribution center based on long-term data of overwintering birds in Finland. Specifically, we explored influences of habitat type, species' snow depth tolerance, species' climatic niche and habitat specialization on range shifts during the past 40 years in 81 bird species. Birds overwintering in arable land shifted more clearly toward north compared to birds of the same species in rural and forest habitats, while the northern range margin shift distances did not significantly differ among the habitat types. Range shifts were more linked with the species' snow depth tolerance rather than species' climatic niche. Snow depth tolerance of species was negatively associated with the eastward shift direction across all habitats, while we found habitat-specific patterns with snow depth for northward shift directions and northern margin shift distances. Species with stronger habitat specializations shifted more strongly toward north as compared to generalist species, whereas the climatic niche of bird species only marginally correlated with range shifts, so that cold-dwelling species shifted longer distances and more clearly eastward. Our study reveals habitat-specific patterns linked to snow conditions for overwintering boreal birds and highlights the importance of habitat availability and preference in climate driven range shifts.

Short-lived species move uphill faster under climate change.

Climate change is pushing species ranges and abundances towards the poles and mountain tops. Although many studies have documented local altitudinal shifts, knowledge of general patterns at a large spatial scale, such as a whole mountain range, is scarce. From a conservation perspective, studying altitudinal shifts in wildlife is relevant because mountain regions often represent biodiversity hotspots and are among the most vulnerable ecosystems. Here, we examine whether altitudinal shifts in birds' abundances have occurred in the Scandinavian mountains over 13 years, and assess whether such shifts are related to species' traits. Using abundance data, we show a clear pattern of uphill shift in the mean altitude of bird abundance across the Scandinavian mountains, with an average speed of 0.9 m per year. Out of 76 species, 7 shifted significantly their abundance uphill. Altitudinal shift was strongly related to species' longevity: short-lived species showed more pronounced uphill shifts in abundance than long-lived species. The observed abundance shifts suggest that uphill shifts are not only driven by a small number of individuals at the range boundaries, but the overall bird abundances are on the move. Overall, the results underscore the wide-ranging impact of climate change and the potential vulnerability of species with slow life histories, as they appear less able to timely respond to rapidly changing climatic conditions.

Covariation in population trends and demography reveals targets for conservation action.

Wildlife conservation policies directed at common and widespread, but declining, species are difficult to design and implement effectively, as multiple environmental changes are likely to contribute to population declines. Conservation actions ultimately aim to influence demographic rates, but targeting actions towards feasible improvements in these is challenging in widespread species with ranges that encompass a wide range of environmental conditions. Across Europe, sharp declines in the abundance of migratory landbirds have driven international calls for action, but actions that could feasibly contribute to population recovery have yet to be identified. Targeted actions to improve conditions on poor-quality sites could be an effective approach, but only if local conditions consistently influence local demography and hence population trends. Using long-term measures of abundance and demography of breeding birds at survey sites across Europe, we show that co-occurring species with differing migration behaviours have similar directions of local population trends and magnitudes of productivity, but not survival rates. Targeted actions to boost local productivity within Europe, alongside large-scale (non-targeted) environmental protection across non-breeding ranges, could therefore help address the urgent need to halt migrant landbird declines. Such demographic routes to recovery are likely to be increasingly needed to address global wildlife declines.

Wintering bird communities are tracking climate change faster than breeding communities.

Global climate change is driving species' distributions towards the poles and mountain tops during both non-breeding and breeding seasons, leading to changes in the composition of natural communities. However, the degree of season differences in climate-driven community shifts has not been thoroughly investigated at large spatial scales. We compared the rates of change in the community composition during both winter (non-breeding season) and summer (breeding) and their relation to temperature changes. Based on continental-scale data from Europe and North America, we examined changes in bird community composition using the community temperature index (CTI) approach and compared the changes with observed regional temperature changes during 1980-2016. CTI increased faster in winter than in summer. This seasonal discrepancy is probably because individuals are less site-faithful in winter, and can more readily shift their wintering sites in response to weather in comparison to the breeding season. Regional long-term changes in community composition were positively associated with regional temperature changes during both seasons, but the pattern was only significant during summer due to high annual variability in winter communities. Annual changes in community composition were positively associated with the annual temperature changes during both seasons. Our results were broadly consistent across continents, suggesting some climate-driven restructuring in both European and North American avian communities. Because community composition has changed much faster during the winter than during the breeding season, it is important to increase our knowledge about climate-driven impacts during the less-studied non-breeding season.

Benefits of protected areas for nonbreeding waterbirds adjusting their distributions under climate warming.

Climate warming is driving changes in species distributions and community composition. Many species have a so-called climatic debt, that is, shifts in range lag behind shifts in temperature isoclines. Inside protected areas (PAs), community changes in response to climate warming can be facilitated by greater colonization rates by warm-dwelling species, but also mitigated by lowering extirpation rates of cold-dwelling species. An evaluation of the relative importance of colonization-extirpation processes is important to inform conservation strategies that aim for both climate debt reduction and species conservation. We assessed the colonization-extirpation dynamics involved in community changes in response to climate inside and outside PAs. To do so, we used 25 years of occurrence data of nonbreeding waterbirds in the western Palearctic (97 species, 7071 sites, 39 countries, 1993-2017). We used a community temperature index (CTI) framework based on species thermal affinities to investigate species turnover induced by temperature increase. We determined whether thermal community adjustment was associated with colonization by warm-dwelling species or extirpation of cold-dwelling species by modeling change in standard deviation of the CTI (CTISD ). Using linear mixed-effects models, we investigated whether communities in PAs had lower climatic debt and different patterns of community change than communities outside PAs. For CTI and CTISD combined, communities inside PAs had more species, higher colonization, lower extirpation, and lower climatic debt (16%) than communities outside PAs. Thus, our results suggest that PAs facilitate 2 independent processes that shape community dynamics and maintain biodiversity. The community adjustment was, however, not sufficiently fast to keep pace with the large temperature increases in the central and northeastern western Palearctic. Our results underline the potential of combining CTI and CTISD metrics to improve understanding of the colonization-extirpation patterns driven by climate warming.

Beneficios de las Áreas Protegidas para las Aves Acuáticas No Reproductoras que Están Ajustando su Distribución Debido al Calentamiento Climático Resumen El calentamiento climático está generando cambios en la distribución y en la composición comunitaria de las especies. Muchas de ellas tienen una deuda climática, es decir, los cambios en la distribución se atrasan con respecto a los cambios en las isoclinas térmicas. Dentro de las áreas protegidas (APs), los cambios comunitarios como respuesta al calentamiento climático pueden facilitarse mediante tasas mayores de colonización por especies de climas cálidos, pero también pueden mitigarse al reducir las tasas de extirpación de las especies de climas fríos. Se requiere una evaluación de la importancia relativa de los procesos de colonización-extirpación para orientar las estrategias de conservación que buscan la reducción de la deuda climática y la conservación de las especies. Analizamos las dinámicas de colonización-extirpación que participan en los cambios comunitarios como respuesta al clima dentro y fuera de las APs. Para realizar lo anterior, usamos datos tomados durante 25 años de la presencia de aves acuáticas no reproductoras en el Paleártico occidental (97 especies, 7,071 sitios, 39 países, 1993-2017). Usamos un marco de trabajo del índice de temperatura comunitaria (ITC) basado en las afinidades térmicas de las especies para así investigar la rotación de especies inducida por el incremento en la temperatura. Determinamos si el ajuste térmico en la comunidad estuvo asociado con la colonización por especies de climas cálidos o con la extirpación de especies de climas fríos al modelar el cambio mediante una desviación estándar del ITC (ITCDS ). Con los modelos lineales de efectos mixtos investigamos si las comunidades dentro de las APs tenían una deuda climática más baja y patrones diferentes de cambio comunitario que las comunidades localizadas fuera de las APs. Con la combinación del ITC y deL ITCDS , las comunidades dentro de las APs tuvieron más especies, una mayor colonización, una menor extirpación y una deuda climática más baja (16%) que las comunidades fuera de las APs. Por lo tanto, nuestros resultados sugieren que las APs facilitan dos procesos independientes que moldean las dinámicas comunitarias y mantienen la biodiversidad. Sin embargo, el ajuste comunitario no fue lo suficientemente rápido para mantener el paso de los grandes incrementos en la temperatura de las regiones central y noreste del Paleártico occidental. Nuestros resultados resaltan el potencial que tiene la combinación de las medidas del ITC y del ICTDS para mejorar el entendimiento de los patrones de colonización-extirpación causados por el calentamiento climático.

The impact of tree crops and temperature on the timing of frugivorous bird migration.

Migration has evolved to tackle temporal changes in availability of resources. Climate change has been shown to affect the migration dates of species, which raises the question of whether the variation in the timing of migration is climate or resource dependent? The relative importance of temperature and availability of food as drivers of migration behaviour during both spring and autumn seasons has been poorly studied. Here, we investigated these patterns in frugivorous and granivorous birds (hereafter frugivorous) that are assumed to postpone their autumn migration when there is plenty of food available, which may also advance upcoming spring migration. On the other hand, especially spring migration dates have been negatively connected with increasing temperatures. We tested whether the autumn and spring migration dates of eleven common frugivorous birds depended on the crop size of trees or ambient temperatures using 29 years of data in Finland. The increased crop sizes of trees delayed autumn migration dates; whereas, autumn temperature did not show a significant connection. We also observed a temporal trend towards later departure. Increasing temperature and crop sizes advanced spring arrival dates. Our results support the hypothesis that the timing of autumn migration in the frugivorous birds depends on the availability of food and is weakly connected with the variation in temperature. Importantly, crop size can have carry-over effects and affect the timing of spring arrival possibly because birds have overwintered closer to the breeding grounds after an abundant crop year.

Protected areas act as a buffer against detrimental effects of climate change-Evidence from large-scale, long-term abundance data.

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

Declining population trends of European mountain birds.

Mountain areas often hold special species communities, and they are high on the list of conservation concern. Global warming and changes in human land use, such as grazing pressure and afforestation, have been suggested to be major threats for biodiversity in the mountain areas, affecting species abundance and causing distribution shifts towards mountaintops. Population shifts towards poles and mountaintops have been documented in several areas, indicating that climate change is one of the key drivers of species' distribution changes. Despite the high conservation concern, relatively little is known about the population trends of species in mountain areas due to low accessibility and difficult working conditions. Thanks to the recent improvement of bird monitoring schemes around Europe, we can here report a first account of population trends of 44 bird species from four major European mountain regions: Fennoscandia, UK upland, south-western (Iberia) and south-central mountains (Alps), covering 12 countries. Overall, the mountain bird species declined significantly (-7%) during 2002-2014, which is similar to the declining rate in common birds in Europe during the same period. Mountain specialists showed a significant -10% decline in population numbers. The slope for mountain generalists was also negative, but not significantly so. The slopes of specialists and generalists did not differ from each other. Fennoscandian and Iberian populations were on average declining, while in United Kingdom and Alps, trends were nonsignificant. Temperature change or migratory behaviour was not significantly associated with regional population trends of species. Alpine habitats are highly vulnerable to climate change, and this is certainly one of the main drivers of mountain bird population trends. However, observed declines can also be partly linked with local land use practices. More efforts should be undertaken to identify the causes of decline and to increase conservation efforts for these populations.

A positive relationship between spring temperature and productivity in 20 songbird species in the boreal zone.

Anthropogenic climate warming has already affected the population dynamics of numerous species and is predicted to do so also in the future. To predict the effects of climate change, it is important to know whether productivity is linked to temperature, and whether species' traits affect responses to climate change. To address these objectives, we analysed monitoring data from the Finnish constant effort site ringing scheme collected in 1987-2013 for 20 common songbird species together with climatic data. Warm spring temperature had a positive linear relationship with productivity across the community of 20 species independent of species' traits (realized thermal niche or migration behaviour), suggesting that even the warmest spring temperatures remained below the thermal optimum for reproduction, possibly due to our boreal study area being closer to the cold edge of all study species' distributions. The result also suggests a lack of mismatch between the timing of breeding and peak availability of invertebrate food of the study species. Productivity was positively related to annual growth rates in long-distance migrants, but not in short-distance migrants. Across the 27-year study period, temporal trends in productivity were mostly absent. The population sizes of species with colder thermal niches had decreasing trends, which were not related to temperature responses or temporal trends in productivity. The positive connection between spring temperature and productivity suggests that climate warming has potential to increase the productivity in bird species in the boreal zone, at least in the short term.

Effects of high latitude protected areas on bird communities under rapid climate change.

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

North by north-west: climate change and directions of density shifts in birds.

There is increasing evidence that climate change shifts species distributions towards poles and mountain tops. However, most studies are based on presence-absence data, and either abundance or the observation effort has rarely been measured. In addition, hardly any studies have investigated the direction of shifts and factors affecting them. Here, we show using count data on a 1000 km south-north gradient in Finland, that between 1970-1989 and 2000-2012, 128 bird species shifted their densities, on average, 37 km towards the north north-east. The species-specific directions of the shifts in density were significantly explained by migration behaviour and habitat type. Although the temperatures have also moved on average towards the north north-east (186 km), the species-specific directions of the shifts in density and temperature did not correlate due to high variation in density shifts. Findings highlight that climate change is unlikely the only driver of the direction of species density shifts, but species-specific characteristics and human land-use practices are also influencing the direction. Furthermore, the alarming results show that former climatic conditions in the north-west corner of Finland have already moved out of the country. This highlights the need for an international approach in research and conservation actions to mitigate the impacts of climate change.

Continent-scale global change attribution in European birds - combining annual and decadal time scales.

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes-based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long-term population growth rates documents simultaneous responses that can be attributed to both climate change and land-use change, including long-term increases in populations of hot-dwelling species and declines in long-distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long-term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high-resolution time series are available in large-scale biodiversity surveys.

Interannual variation and long-term trends in proportions of resident individuals in partially migratory birds.

Partial migration - a part of a population migrates and another part stays resident year-round on the breeding site - is probably the most common type of migration in the animal kingdom, yet it has only lately garnered more attention. Theoretical studies indicate that in partially migratory populations, the proportion of resident individuals (PoR) should increase in high latitudes in response to the warming climate, but empirical evidence exists for few species. We provide the first comprehensive overview of the environmental factors affecting PoR and the long-term trends in PoR by studying 27 common partially migratory bird species in Finland. The annual PoR values were calculated by dividing the winter bird abundance by the preceding breeding abundance. First, we analysed whether early-winter temperature, winter temperature year before or the abundance of tree seeds just before overwintering explains the interannual variation in PoR. Secondly, we analysed the trends in PoR between 1987 and 2011. Early-winter temperature explained the interannual variation in PoR in the waterbirds (waterfowl and gulls), most likely because the temperature affects the ice conditions and thereby the feeding opportunities for the waterbirds. In terrestrial species, the abundance of seeds was the best explanatory variable. Previous winter's temperature did not explain PoR in any species, and thus, we conclude that the variation in food availability caused the interannual variation in PoR. During the study period, PoR increased in waterbirds, but did not change in terrestrial birds. Partially migratory species living in physically contrasting habitats can differ in their annual and long-term population-level behavioural responses to warming climate, possibly because warm winter temperatures reduce ice cover and improve the feeding possibilities of waterbirds but do not directly regulate the food availability for terrestrial birds.

Velocity of density shifts in Finnish landbird species depends on their migration ecology and body mass.

A multitude of studies confirm that species have changed their distribution ranges towards higher elevations and towards the poles, as has been predicted by climate change forecasts. However, there is large interspecific variation in the velocity of range shifts. From a conservation perspective, it is important to understand which factors explain variation in the speed and the extent of range shifts, as these might be related to the species' extinction risk. Here, we study shifts in the mean latitude of occurrence, as weighted by population density, in different groups of landbirds using 40 years of line transect data from Finland. Our results show that the velocity of such density shifts differed among migration strategies and increased with decreasing body size of species, while breeding habitat had no influence. The slower velocity of large species could be related to their longer generation time and lower per capita reproduction that can decrease the dispersal ability compared to smaller species. In contrast to some earlier studies of range margin shifts, resident birds and partial migrants showed faster range shifts, while fully migratory species were moving more slowly. The results suggest that migratory species, especially long-distance migrants, which often show decreasing population trends, might also have problems in adjusting their distribution ranges to keep pace with global warming.