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.

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.

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.

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.

The potential for dispersal of microalgal resting cysts by migratory birds.

Most microalgal species are geographically widespread, but little is known about how they are dispersed. One potential mechanism for long-distance dispersal is through birds, which may transport cells internally (endozoochory) and deposit them during, or in-between, their migratory stopovers. We hypothesize that dinoflagellates, in particular resting stages, can tolerate bird digestion; that bird temperature, acidity, and retention time negatively affect dinoflagellate viability; and that recovered cysts can germinate after passage through the birds' gut, contributing to species-specific dispersal of the dinoflagellates across scales. Tolerance of two dinoflagellate species (Peridiniopsis borgei, a warm-water species and Apocalathium malmogiense, a cold-water species) to Mallard gut passage was investigated using in vitro experiments simulating the gizzard and caeca conditions. The effect of in vitro digestion and retention time on cell integrity, cell viability, and germination capacity of the dinoflagellate species was examined targeting both their vegetative and resting stages. Resting stages (cysts) of both species were able to survive simulated bird gut passage, even if their survival rate and germination were negatively affected by exposure to acidic condition and bird internal temperature. Cysts of A. malmogiense were more sensitive than P. borgei to treatments and to the presence of digestive enzymes. Vegetative cells did not survive conditions of bird internal temperature and formed pellicle cysts when exposed to gizzard-like acid conditions. We show that dinoflagellate resting cysts serve as dispersal propagules through migratory birds. Assuming a retention time of viable cysts of 2-12 h to duck stomach conditions, cysts could be dispersed 150-800 km and beyond.

Community occupancy before-after-control-impact (CO-BACI) analysis of Hurricane Gudrun on Swedish forest birds.

Resilience of ecological communities to perturbation is important in the face of increased global change from anthropogenic stressors. Monitoring is required to detect the impact of, and recovery from, perturbations, and before-after-control-impact (BACI) analysis provides a powerful framework in this regard. However, species in a community are not observed with perfect detection, and occupancy analysis is required to correct for imperfect detectability of species. We present a Bayesian community occupancy before-after-control-impact (CO-BACI) framework to monitor ecological community response to perturbation when constituent species are imperfectly detected. We test the power of the model to detect changes in community composition following an acute perturbation with simulation. We then apply the model to a study of the impact of a large hurricane on the forest bird community of Sweden, using data from the national bird survey scheme. Although simulation shows the model can detect changes in community occupancy following an acute perturbation, application to a Swedish forest bird community following a major hurricane detected no change in community occupancy despite widespread forest loss. Birds with landscape occupancy less than 50% required correcting for detectability. We conclude that CO-BACI analysis is a useful tool that can incorporate rare species in analyses and detect occupancy changes in ecological communities following perturbation, but, because it does not include abundance, some impacts may be overlooked.

Impact of climate change on communities: revealing species' contribution.

1. Although climate is known to play an important role in structuring biological communities, high-resolution analyses of recent climatic impacts on multiple components of diversity are still sparse. Additionally, there is a lack of knowledge about which species drive community response to environmental change. 2. We used a long-term breeding bird data set that encompasses a large latitudinal and altitudinal range to model the effect of temperature on spatial and temporal patterns in alpha and beta diversity. We also established a novel framework for identifying species-specific contributions to these macroecological patterns, hence combining two different approaches for identifying climatic impacts. 3. Alpha diversity increased over time, whilst beta diversity declined; both diversity metrics showed a significant relationship with recent temperature anomalies. By partitioning beta diversity, we showed that the decline was predominately driven by changes in species turnover rather than nestedness suggesting a process of replacement by more common species. 4. Using jackknife analyses we identified how individual species influenced the modelled relationships of diversity with temperature and time. Influential species tended to be habitat generalists with moderate to large distributions. 5. We demonstrate that different facets of avian diversity can respond rapidly to temperature anomalies and as a result have undergone significant changes in the last decade. In general, it appears that warming temperatures are driving compositional homogenization of temperate bird communities via range expansion of common generalist species.

Åke Lindström.

Q&A with Åke Lindström who studies bird migration and biology.

Solar heating may explain extreme diel flight altitude changes in migrating birds.

Great reed warblers, Acrocephalus arundinaceus,1 and great snipes, Gallinago media,2 exhibit a diel cycle in flight altitudes-flying much higher during the day than the night-when performing migratory flights covering both night and day. One hypothesis proposed to explain this behavior is that the birds face additional heating by solar radiation during daytime and hence must climb to very high, and thus also very cold, altitudes to avoid overheating during daytime flights.1,2 Yet, solar heat gain in birds has been shown to drastically decrease with wind speed,3,4 and the quantitative heating effect by solar radiation on a bird flying with an airspeed of 10 m/s or more is unknown. We analyzed temperature data from multisensor data loggers (MDLs)5,6 placed without direct exposure to solar radiation on great reed warblers (the logger covered by feathers on the back) and great snipes (the logger on the leg, covered from the sun by the tail). We found that logger temperatures were significantly higher (5.9°C-8.8°C in great reed warblers and 4.8°C-5.4°C in great snipes) during the day than during the night in birds flying at the same altitudes (and thus also the same expected ambient air temperatures). These results strongly indicate that the heat balance of the flying birds is indeed affected by solar radiation, which is in accordance with the hypothesis that solar radiation is a key factor causing the remarkable diel cycles in flight altitude observed in these two long-distance migrant bird species.1,2.

Local colonisations and extinctions of European birds are poorly explained by changes in climate suitability.

Climate change has been associated with both latitudinal and elevational shifts in species' ranges. The extent, however, to which climate change has driven recent range shifts alongside other putative drivers remains uncertain. Here, we use the changing distributions of 378 European breeding bird species over 30 years to explore the putative drivers of recent range dynamics, considering the effects of climate, land cover, other environmental variables, and species' traits on the probability of local colonisation and extinction. On average, species shifted their ranges by 2.4 km/year. These shifts, however, were significantly different from expectations due to changing climate and land cover. We found that local colonisation and extinction events were influenced primarily by initial climate conditions and by species' range traits. By contrast, changes in climate suitability over the period were less important. This highlights the limitations of using only climate and land cover when projecting future changes in species' ranges and emphasises the need for integrative, multi-predictor approaches for more robust forecasting.

More and more generalists: two decades of changes in the European avifauna.

Biotic homogenization (BH) is a process whereby some species (losers) are systematically replaced by others (winners). While this process has been related to the effects of anthropogenic activities, whether and how BH is occurring across regions and the role of native species as a driver of BH has hardly been investigated. Here, we examine the trend in the community specialization index (CSI) for 234 native species of breeding birds at 10,111 sites in six European countries from 1990 to 2008. Unlike many BH studies, CSI uses abundance information to estimate the balance between generalist and specialist species in local assemblages. We show that bird communities are more and more composed of native generalist species across regions, revealing a strong, ongoing BH process. Our result suggests a rapid and non-random change in community composition at a continental scale is occurring, most likely driven by anthropogenic activities.

Great flights by great snipes: long and fast non-stop migration over benign habitats.

Migratory land birds perform extreme endurance flights when crossing ecological barriers, such as deserts, oceans and ice-caps. When travelling over benign areas, birds are expected to migrate by shorter flight steps, since carrying the heavy fuel loads needed for long non-stop flights comes at considerable cost. Here, we show that great snipes Gallinago media made long and fast non-stop flights (4300-6800 km in 48-96 h), not only over deserts and seas but also over wide areas of suitable habitats, which represents a previously unknown migration strategy among land birds. Furthermore, the great snipes achieved very high ground speeds (15-27 m s(-1)), which was not an effect of strong tailwind support, and we know of no other animal that travels this rapidly over such a long distance. Our results demonstrate that some migratory birds are prepared to accept extreme costs of strenuous exercise and large fuel loads, even when stopover sites are available along the route and there is little tailwind assistance. A strategy of storing a lot of energy before departure, even if migration is over benign habitats, may be advantageous owing to differential conditions of fuel deposition, predation or infection risk along the migration route.

Extreme altitude changes between night and day during marathon flights of great snipes.

Several factors affect the flight altitude of migratory birds, such as topography, ambient temperature, wind conditions, air humidity, predation avoidance, landmark orientation, and avoiding over-heating from direct sunlight.1-6 Recent tracking of migratory birds over long distances has shown that migrants change flight altitude more commonly and dramatically than previously thought.4-8 The reasons behind these altitude changes are not well understood. In their seasonal migrations between Sweden and sub-Saharan Africa, great snipes Gallinago media make non-stop flights of 4,000-7,000 km, lasting 60-90 h.9,10 Activity and air pressure data from multisensor dataloggers showed that great snipes repeatedly changed altitudes around dawn and dusk, between average cruising heights about 2,000 m (above sea level) at night and around 4,000 m during daytime. Frequency and autocorrelation analyses corroborated a conspicuous diel cycle in flight altitude. Most birds regularly flew at 6,000 m and one bird reached 8,700 m, possibly the highest altitude ever recorded for an identified migrating bird. The diel altitude changes took place independently of climate zone, topography, and habitat overflown. Ambient temperature, wind condition, and humidity have no important diel variation at the high altitudes chosen by great snipes. Instead, improved view for orientation by landmarks, predator avoidance, and not least, seeking cold altitudes at day to counteract heating from direct sunlight are the most plausible explanations for the diel altitude cycle. Together with similar recent findings for a small songbird,6 the great snipes' altitudinal performance sheds new light on the complexity and challenges of migratory flights.

Long-term and large-scale multispecies dataset tracking population changes of common European breeding birds.

Around fifteen thousand fieldworkers annually count breeding birds using standardized protocols in 28 European countries. The observations are collected by using country-specific and standardized protocols, validated, summarized and finally used for the production of continent-wide annual and long-term indices of population size changes of 170 species. Here, we present the database and provide a detailed summary of the methodology used for fieldwork and calculation of the relative population size change estimates. We also provide a brief overview of how the data are used in research, conservation and policy. We believe this unique database, based on decades of bird monitoring alongside the comprehensive summary of its methodology, will facilitate and encourage further use of the Pan-European Common Bird Monitoring Scheme results.

Bird population trends are linearly affected by climate change along species thermal ranges.

Beyond the effects of temperature increase on local population trends and on species distribution shifts, how populations of a given species are affected by climate change along a species range is still unclear. We tested whether and how species responses to climate change are related to the populations locations within the species thermal range. We compared the average 20 year growth rates of 62 terrestrial breeding birds in three European countries along the latitudinal gradient of the species ranges. After controlling for factors already reported to affect bird population trends (habitat specialization, migration distance and body mass), we found that populations breeding close to the species thermal maximum have lower growth rates than those in other parts of the thermal range, while those breeding close to the species thermal minimum have higher growth rates. These results were maintained even after having controlled for the effect of latitude per se. Therefore, the results cannot solely be explained by latitudinal clines linked to the geographical structure in local spring warming. Indeed, we found that populations are not just responding to changes in temperature at the hottest and coolest parts of the species range, but that they show a linear graded response across their European thermal range. We thus provide insights into how populations respond to climate changes. We suggest that projections of future species distributions, and also management options and conservation assessments, cannot be based on the assumption of a uniform response to climate change across a species range or at range edges only.

Consequences of organic farming and landscape heterogeneity for species richness and abundance of farmland birds.

It has been suggested that organic farming may benefit farmland biodiversity more in landscapes that have lost a significant part of its former landscape heterogeneity. We tested this hypothesis by comparing bird species richness and abundance during the breeding season in organic and conventional farms, matched to eliminate all differences not directly linked to the farming practice, situated in either homogeneous plains with only a little semi-natural habitat or in heterogeneous farmland landscapes with abundant field borders and semi-natural grasslands. The effect of farm management on species richness interacted with landscape structure, such that there was a positive relationship between organic farming and diversity only in homogeneous landscapes. This pattern was mainly dependent on the species richness of passerine birds, in particular those that were invertebrate feeders. Species richness of non-passerines was positively related to organic farming independent of the landscape context. Bird abundance was positively related to landscape heterogeneity but not to farm management. This was mainly because the abundance of passerines, particularly invertebrate feeders, was positively related to landscape heterogeneity. We suggest that invertebrate feeders particularly benefit from organic farming because of improved foraging conditions through increased invertebrate abundances in otherwise depauperate homogeneous landscapes. Although many seed-eaters also benefit from increased insect abundance, they may also utilize crop seed resources in homogeneous landscapes and conventional farms. The occurrence of an interactive effect of organic farming and landscape heterogeneity on bird diversity will have consequences for the optimal allocation of resources to restore the diversity of farmland birds.

Are birds stressed during long-term flights? A wind-tunnel study on circulating corticosterone in the red knot.

During endurance flight most birds do not feed and have to rely on their body reserves. Fat and protein is catabolised to meet the high energetic demands. Even though the hormonal regulation of migration is complex and not yet fully understood, the adrenocortical hormone corticosterone crystallizes to play a major role in controlling physiological traits in migratory birds during flight. However, results from field studies are partially equivocal, not least because data from birds during endurance flight are hard to get and present mostly a momentary shot. A wind-tunnel experiment offered the possibility to measure repeatedly under controlled conditions the effect of long flights on the stress hormone corticosterone. In a long-distance migrating shorebird, the red knot Calidris canutus, we measured plasma concentrations of corticosterone within 3 min and after a restraint time of 30 min directly after 2h and 10h non-stop flights, respectively, and during rest. Baseline corticosterone levels were unchanged directly after the flights, indicating that endurance flight did not affect corticosterone levels. The adrenocortical response to restraint showed the typical rise in birds during rest, while birds after a 2 or 10h flight substantially decreased plasma corticosterone concentrations. We suggest that the negative adrenocortical response to restraint after flight is part of the mechanism to reduce the proteolytic effect of corticosterone to save muscle protein and to avoid muscle damaging effects.