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Aim: Species distribution models (SDMs) are widely used to make predictions on how species distributions may change as a response to climatic change. To assess the reliability of those predictions, they need to be critically validated with respect to what they are used for. While ecologists are typically interested in how and where distributions will change, we argue that SDMs have seldom been evaluated in terms of their capacity to predict such change. Instead, typical retrospective validation methods estimate model's ability to predict to only one static time in future. Here, we apply two validation methods, one that predicts and evaluates a static pattern, while the other measures change and compare their estimates of predictive performance. Location: Fennoscandia.Methods: We applied a joint SDM to model the distributions of 120 bird species in four model validation settings. We trained models with a dataset from 1975 to 1999 and predicted species' future occurrence and abundance in two ways: for one static time period (2013- 2016, "static validation') and for a change between two time periods (difference between 1996- 1999 and 2013- 2016, "change validation'). We then measured predictive performance using correlation between predicted and observed values. We also related predictive performance to species traits. Results: Even though static validation method evaluated predictive performance as good, change method indicated very poor performance. Predictive performance was not strongly related to any trait.Main Conclusions: Static validation method might overestimate predictive performance by not revealing the model's inability to predict change events. If species' distributions remain mostly stable, then even an unfit model can predict the near future well due to temporal autocorrelation. We urge caution when working with forecasts of changes in spatial patterns of species occupancy or abundance, even for SDMs that are based on time series datasets unless they are critically validated for forecasting such change.

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.

There is presently great concern about local, regional and global loss of species. We examined those breeding species and subspecies (“species”) of birds that either established themselves or went extinct in Sweden in 1850–2009. In this period Sweden got 38 new and lost twelve breeding bird species. The average (and surprisingly steady) rate of colonization was 2.4 species per decade, which was three times higher than the rate of extinction of 0.75 species per decade. We also predict future establishments and extinctions until year 2050, based on Species Distribution Modelling and recent information from neighbouring countries. We find it probable that eleven new species will join the Swedish avifauna until 2050 and possible that another nine species will do so. The predicted rate of probable establishment is 2.9 species per decade. We identified seven species/subspecies to be at risk of extinction by 2050. The future in Sweden for several of these species is probably dependent on the success or failure of ongoing conservation programs. There seems to be no immediate threat of establishment of alien species.

Climate change is pushing species ranges towards 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 very limited. From a conservation perspective, studying altitudinal shifts is particularly important as mountain regions often represent biodiversity hotspots and are among the most vulnerable ecosystems. Here, we examine whether altitudinal shifts have occurred among birds 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 shifts in the mean altitudes of the bird species’ abundances across the Scandinavian mountains, with an average speed of 0.9 m per year. Out of 77 species, 54 shifted their ranges uphill. In general, the range shift was faster when the altitudinal range within the area was wider. Importantly, the altitudinal shift was strongly related to species’ longevity: short-lived species showed more pronounced altitudinal uphill shifts than long-lived species. Our results show that the altitudinal range shifts are not only driven by a small number of individuals at the range boundaries, but the overall bird abundances are on the move. This highlights the wide-ranging impact of climate change and the potential vulnerability of species with slow life-histories, as they appear unable to timely respond to rapidly changing climatic conditions.

Many bird species typical to farmland have declined in numbers in recent decades. Some farmland bird species occur outside farmland in other open man-made habitats, such as forest clear-cuts. Given that in Sweden the total area of clear-cuts (5–6 %) almost equals that of farmland (8 %), clear-cuts have the potential to be an important habitat for some farmland birds. We investigated how bird community composition and species abundance on Swedish clear-cuts varied with geographical region, clear-cut characteristics and the surrounding landscape, with special focus on farmland species. Based on short visits in 2017–2019, we surveyed the occurrence of birds on 300 representative clear-cuts in Sweden, including six regions and 1300 km of latitude. We counted 1170 birds of 66 different species, including 10 out of 15 bird species in existing farmland bird indicators. Both bird community composition and species abundance varied with clear-cut size, age, vegetation height, region and proportion of nearby farmland, depending on species’ ecology. All six farmland species being proportionally common on clear-cuts are, when in farmland, dependent on residual habitats such as pastures and field borders with bushes and trees. In contrast, a third of the dedicated farmland species were scarce in clear-cuts, probably avoiding them because of predator-related avoidance of forest edges and/or a lack of suitable nest sites or food resources. To determine the full importance of forest clear-cuts for farmland species, and potentially make them more favourable for farmland birds, more studies are needed on the reproductive success on clear-cuts and which site features make them suitable habitat. This study did not determine the consequences of clear-cuts for forest birds across the forest cycle, consequences which needs to be balanced against any gains for farmland birds.

Funding: The Swedish Bird Survey was supported by grants from the Swedish Environmental Protection Agency, with additional financial and logistic support from the Regional County Boards (Länsstyrelsen). The surveys were carried out within the framework of the strategic research environment Biodiversity and Ecosystem Services in a Changing Climate (BECC). The Norwegian breeding bird monitoring (Norsk hekkefuglovervåking) was financed by the Norwegian Environment Agency and the Ministry of Climate and Environment. Our research was funded through the 2017-2018 Belmont Forum and BiodivERsA joint call for research proposals, under the BiodivScen ERA-Net COFUND programme with financial support from the Research Council of Norway (295767), the Swedish Research Council FORMAS (2018-02440, 2018-02441), the Academy of Finland (323527, 326327, 326338) and the U.S. National Science Foundation (ICER-1927646). The relationships between species abundance or occurrence versus spatial variation in climate are commonly used in species distribution models (SDMs) 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 spatio-temporal 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 modelled 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 spatio-temporal 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. Publisher PDF

Corrigendum: Ecological Indicators, Volume 137, April 2022, Article number 108627, https://doi.org/10.1016/j.ecolind.2022.108627. In order to understand species' sensitivity to habitat change, we must correctly determine if a species is associated with a habitat or not, and if it is associated, its degree of specialization for that habitat. However, definitions of species' habitat association and specialization are often static, categorical classifications that coarsely define species as either habitat specialists or generalists and can fail to account for potential temporal or spatial differences in association or specialization. In contrast, quantitative metrics can provide a more nuanced assessment, defining species' habitat associations and specialization along a continuous scale and accommodate for temporal or spatial variation, but these approaches are less widely used. Here we explore relative habitat use (RHU) as a metric for quantifying species' association with and degree of specialization for different habitat types. RHU determines the extent of a species' association with a given habitat by comparing its abundance in that habitat relative to its mean abundance across all other habitats. Using monitoring data for breeding birds across Europe from 1998 to 2017; we calculate RHU scores for 246 species for five habitat types and compared them to the literature-based classifications of their association with and specialization for each of these habitats. We also explored the temporal variation in species' RHU scores for each habitat and assessed how this varied according to association and degree of specialization. In general, species' RHU and literature-derived classifications were well aligned, as RHU scores for a given habitat increased in line with reported association and specialization. In addition, temporal variation in RHU scores were influenced by association and degree of specialization, with lower scores for those associated with, and those more specialized to, a given habitat. As a continuous metric, RHU allows a detailed assessment of species' association with and degree of specialization for different habitats that can be tailored to specific temporal and/or spatial requirements. It has the potential to be a valuable tool for identifying indicator species and in supporting the design, implementation and monitoring of conservation management actions.

Natural sounds, and bird song in particular, play a key role in building and maintaining our connection with nature, but widespread declines in bird populations mean that the acoustic properties of natural soundscapes may be changing. Using data-driven reconstructions of soundscapes in lieu of historical recordings, here we quantify changes in soundscape characteristics at more than 200,000 sites across North America and Europe. We integrate citizen science bird monitoring data with recordings of individual species to reveal a pervasive loss of acoustic diversity and intensity of soundscapes across both continents over the past 25 years, driven by changes in species richness and abundance. These results suggest that one of the fundamental pathways through which humans engage with nature is in chronic decline, with potentially widespread implications for human health and well-being., Birdsong has long connected humans to nature. Historical reconstructions using bird monitoring and song recordings collected by citizen scientists reveal that the soundscape of birdsong in North America and Europe is both quieter and less varied, mirroring declines in bird diversity and abundance.

Waders form a conspicuous part of the bird fauna in boreal and arctic areas, where they inhabit forests, wetlands, mires and tundra. These are important breeding areas for a large set of wader species, and may be particularly vulnerable to climate change. However, large-scale and systematic monitoring data from the breeding grounds of boreal and arctic waders are largely lacking. We present population trends for 22 wader species breeding in the boreal and arctic parts of Fennoscandia (Norway, Sweden and Finland) between 2006 and 2018. The trends are based on 9,713 surveys of 1,505 unique routes (6–8 km), each surveyed in at least two years, evenly distributed over an area of ~1 million km2. The trends were significantly negative for three species: Red-necked Phalarope Phalaropus lobatus (–7.9% year-1), Broad-billed Sandpiper Calidris falcinellus (–5.4% year-1), and Whimbrel Numenius phaeopus (–1.3% year-1). The trends were significantly positive for three species: Oystercatcher Haematopus ostralegus (+4.9% year-1), Dunlin Calidris a. alpina (+4.2% year-1) and Wood Sandpiper Tringa glareola (+0.8% year-1). For the remaining species, we found no statistically significant trends. On average, as shown by a multi-species indicator, there was no general change in numbers over time. On 1,539 routes with at least one survey, wader species richness as well as total number of wader pairs increased significantly with increasing latitude. Species population trend was not correlated with breeding latitude, but population trends of long-distance migrants tended to be more negative than those of medium-distance migrants. The recent fortunes of waders breeding in northern Fennoscandia have been more buoyant than those in other parts of Europe, but the trends for some species are worrying. © 2019, International Wader Study Group. All rights reserved.

The effect that extreme natural events have on biological diversity is relatively poorly known. We used a before–after control-impact (BACI) design to analyze changes in bird abundances and communities following Hurricane Gudrun, which struck southern Sweden in January 2005, felling 75 million m3 of forest and causing damage to 5% of forested areas (half a million hectares) in a few hours. We used recent measures of impact in combination with classical BACI contrasts to analyze bird count data from a monitoring program in Sweden. We investigated changes in the abundance of 17 species commonly found in forests, along with changes in species composition and functional structure of the bird community. In total, we considered 34 response variables and examined whether responses were immediate or long-term. There was no evidence of a strong effect of the hurricane on the abundances of six species. Estimates of the effects on five species were too uncertain to draw inferences. We detected positive and negative effects of the hurricane on the abundances of the remaining six species, but the magnitude of effects often was small. Generally, the effects were in the expected direction: negative on birds associated with mature forest and positive on birds associated with open land or young forest. We found evidence of an increase in the proportion of species that nest on the ground and a decrease in the proportion of species that nest in cavities and trees. In contrast, the hurricane had no discernible effect on functional measures of diversity (richness, evenness or divergence), or on communities’ reproductive or morphological characteristics. Our results suggest that the hurricane affected bird populations and communities, but the magnitude of effects was generally small. (Less)

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.

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 , 2 , 3 , 4 , 5 , 6 Recent tracking of migratory birds over long distances has shown that migrants change flight altitude more commonly and dramatically than previously thought. 4 , 5 , 6 , 7 , 8 The reasons behind these altitude changes are not well understood. In theirseasonal 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.

Goose management in Europe is faced by multiple challenges, as some species are declining and in need of conservation actions, while other populations have become very abundant, resulting in calls for increased harvest. Sweden has long-Term series of harvest data and counts of breeding and autumn-staging geese. We used national data (indices) for greylag goose, bean goose and Canada goose to study shifts in temporal trends and correlative patterns, and to infer possible causal links between harvest and population trends. Our study provides an opportunity to guide management given the data collected within the present monitoring, as well as to suggest improvements for future data collection. The populations of greylag and Canada geese increased in Sweden 1979-2018, but this long-Term trend included a recent decrease in the latter species. Bean goose breeding index decreased, whilst staging numbers and harvest varied with no clear long-Term trend. For Canada goose, our analysis suggests that harvest may affect population growth negatively. For bean goose and greylag goose we could not detect any effect of harvest on autumn counts the following year. We find that the present data and analysis of coherence may suffice as basis for decisions for the current management situation in Sweden with its rather unspecific goals for greylag (very abundant) and Canada goose (invasive species) populations. However, for management of bean geese, with international concerns of over harvest, data lack crucial information. For future management challenges, with more explicit goals, for all goose species we advocate information that is more precise. Data such as hunting effort, age-structure of goose populations and mark-recapture data to estimate survival and population size, is needed to feed predictive population models guiding future Swedish and European goose management. (Less)

Agricultural landscapes have changed fast in Europe, which has led to steep declines in farmland biodiversity. While this has differentially impacted “farmland species”, some of these seem to be able to use alternative man-made open habitats in forests, such as forest clear-cuts and powerline rights-of-way. We investigated the use of clear-cuts by two historically declining farmland birds, the Yellowhammer (Emberiza citrinella) and the Red-backed Shrike (Lanius collurio), to determine both local and landscape habitat characteristics of clear-cuts used for breeding. Among 101 clear-cuts visited in Scania, southern Sweden, 27% were occupied by Yellowhammers and 12% by Red-backed Shrikes. Yellowhammer occurrence in clear-cuts was positively related to local spruce cover (the planted tree species). There was also an interaction between clear-cut size and the proportion of farmland in the landscape, such that the positive effect of the proportion of farmland nearby was stronger for larger clear-cuts. Red-backed Shrike occurrence was positively related to clear-cut size and the local presence of piles of forest residues (tops and branches). The relation between occurrence and the proportion of clear-cuts in the landscape was positive for small clear-cuts and negative for larger ones. Yellowhammers and Red-backed shrikes are affected by both the local habitat structure in the clearcut and on the composition of the surrounding landscape, most likely because they use the clear-cut for nesting and the surroundings as complementary feeding habitat. To what extent clear-cuts as complementary breeding habitat to farmland may contribute to the conservation of Yellowhammers and Red-backed shrikes, as well as other farmland birds, is a promising path for future investigations.

Waders normally fly with their legs stretched out under the tail, with their feet pointing backwards. Whereas there are a few published records of waders flying with their legs folded forward (like a passerine), with legs and feet tucked in under the belly feathers, the causes and consequences of waders flying with folded legs are poorly understood. Through searching the literature, asking ornithologists in person and through public calls, and from searching photos on the internet, we retrieved 90 records of 17 different species where one or more individuals flew with folded legs. It was most common in Spotted Redshank Tringa erythropus (in up to 10% of all birds) and Common Redshank T. totanus, but was also observed in short-legged species such as Dunlin Calidris alpina and Common Ringed Plover Charadrius hiaticula. The behaviour was recorded in most months of the year, in different age classes, in single birds as well as in flocks, at different ambient temperatures, and in both migratory flights and local move-ments. Whereas reducing heat loss at low temperatures is a likely, but not omnipo-tent, explanation for waders flying with folded legs, aerodynamics and leg muscle fatigue may also affect how waders keep their legs in flight. (Less)

The European Union's Natura 2000 (N2000) is among the largest international networks of protected areas. One of its aims is to secure the status of a predetermined set of (targeted) bird and butterfly species. However, nontarget species may also benefit from N2000. We evaluated how the terrestrial component of this network affects the abundance of nontargeted, more common bird and butterfly species based on data from long-term volunteer-based monitoring programs in 9602 sites for birds and 2001 sites for butterflies. In almost half of the 155 bird species assessed, and particularly among woodland specialists, abundance increased (slope estimates ranged from 0.101 [SD 0.042] to 3.51 [SD 1.30]) as the proportion of landscape covered by N2000 sites increased. This positive relationship existed for 27 of the 104 butterfly species (estimates ranged from 0.382 [SD 0.163] to 4.28 [SD 0.768]), although most butterflies were generalists. For most species, when land-cover covariates were accounted for these positive relationships were not evident, meaning land cover may be a determinant of positive effects of the N2000 network. The increase in abundance as N2000 coverage increased correlated with the specialization index for birds, but not for butterflies. Although the N2000 network supports high abundance of a large spectrum of species, the low number of specialist butterflies with a positive association with the N2000 network shows the need to improve the habitat quality of N2000 sites that could harbor open-land butterfly specialists. For a better understanding of the processes involved, we advocate for standardized collection of data at N2000 sites.Efectos de Natura 2000 sobre las Especies No Focales de Aves y Mariposas con Base en Datos de Ciencia Ciudadana Resumen La red Natura 2000 (N2000) de la Unión Europea está entre las redes internacionales más grandes de áreas protegidas. Uno de sus objetivos es asegurar el estado de un conjunto predeterminado de especies de aves y mariposas (focales). Sin embargo, las especies no focales también pueden beneficiarse con la N2000. Evaluamos cómo el componente terrestre de esta red afecta la abundancia de las especies de aves y mariposas no focales más comunes con base en los datos de programas de monitoreo voluntario a largo plazo en 9,602 sitios para aves y en 2,001 sitios para mariposas. En casi la mitad de las 155 especies de aves evaluadas, particularmente entre aquellas especies especialistas en zonas boscosas, la abundancia incrementó (los estimaciones de la pendiente variaron desde 0.101 [DS 0.042] hasta 3.51 [DS 1.30]) conforme incrementó la proporción del paisaje cubierto por sitios de la N2000. Esta relación positiva existió en 27 de las 104 especies de mariposas (con una variación de estimaciones desde 0.382 [DS 0.163] hasta 4.28 [DS 0.768]), aunque la mayoría de las especies de mariposas fueron generalistas. Cuando se consideraron las covarianzas de cobertura de suelo estas relaciones positivas no fueron evidentes para la mayoría de las especies, lo que significa que la cobertura de suelo puede ser una determinante de los efectos positivos de la red N2000. El incremento en la abundancia conforme aumentó la cobertura de la N2000 estuvo correlacionado con el índice de especialización de las aves, pero no el de las mariposas. Aunque la red N2000 sostiene la abundancia alta de un espectro amplio de especies, el bajo número de mariposas especialistas con una asociación positiva a la red N2000 demuestra la necesidad de mejorar la calidad del hábitat de los sitios N2000 que podrían albergar a mariposas especialistas de campo abierto. Para un mejor entendimiento de los procesos involucrados, promovemos una recolección estandarizada de datos en los sitios de la red N2000.欧盟的 Natura 2000(N2000) 是全球最大的保护地网络之一, 它的目标之一是保护一批目标鸟类和蝴蝶物种的现状。不过, 非目标物种也可能从 N2000 中获益。本研究评估了该网络陆地保护区如何影响更为常见的非目标鸟类和蝴蝶的丰度, 所用数据来自长期的志愿者监测项目, 包括了 9602 个鸟类监测点和 2001 个蝴蝶监测点。在评估的 155 种鸟类中, 几乎一半的鸟类, 特别是林地专性种, 在 N2000 位点覆盖的景观比例上升时数量增加 (斜率估计值在 0.101 [SD 0.042] 到3.51 [SD 1.30] 之间) 。虽然大多数蝴蝶都是广幅种, 但 104 种蝴蝶中也有 27 种存在这种正相关关系 (斜率估计值在 0.382 [SD 0.163] 到 4.28 [SD 0.768]) 。对于大多数物种来说, 当考虑土地覆盖协变量时, 这样的正相关关系并不明显, 这意味着土地覆盖可能是 N2000 网络能否对物种产生积极影响的关键因素。物种的丰度随着 N2000 覆盖度增加而增加的现象与鸟类的生境特化指数相关, 而在蝴蝶中则无关。虽然 N2000 网络使许多物种得以维持较高丰度, 但获得其积极影响的蝴蝶专性种数量仍较少, 这说明需要在 N2000 网络中提高专性生活在开阔地的蝴蝶的生境质量。为了更好地理解其中的过程, 我们建议收集更多标准化的 N2000 位点数据。 【翻译: 胡怡思; 审校: 聂永刚】.

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Many bird populations in temperate regions have advanced their timing of breeding in response to a warming climate in recent decades. However, long-term trends in temperature differ geographically and between seasons, and so do responses of local breeding populations. Data on breeding bird phenology from subarctic and arctic passerine populations are scarce, and relatively little data has been recorded in open-nesting species. We investigated the timing of breeding and its relationship to spring temperature of 14 mainly open-nesting passerine species in subarctic Swedish Lapland over a period of 32 years (1984–2015). We estimated timing of breeding from the progress of post-juvenile moult in mist-netted birds, a new method exploring the fact that the progress of post-juvenile moult correlates with age. Although there was a numerical tendency for earlier breeding in most species (on average −0.09 days/year), changes were statistically significant in only three species (by −0.16 to −0.23 days/year). These figures are relatively low compared with what has been found in other long-term studies but are similar to a few other studies in subarctic areas. Generally, annual hatching dates were negatively correlated with mean temperature in May. This correlation was stronger in long-distance than in short-distance migrants. Although annual temperatures at high northern latitudes have increased over recent decades, there was no long-term increase in mean temperature in May over the study period at this subarctic site. This is probably the main reason why there were only small long-term changes in hatching dates. (Less)

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.

Northern European peatlands are important habitats for biological conservation because they support rich biodiversity and unique species compositions. However, historical management of peatland habitats has had negative consequences for biodiversity and their degradation remains a major conservation concern. Despite increasing awareness of the conservation value of peatlands, the statuses and ecological requirements of peat land species have remained largely understudied. Here, we first analysed temporal trends of Northern European peatland birds to document the status of their populations using bird data from five different countries. Second, we used Finnish monitoring data to assess habitat preferences of peatland bird species, hence helping to target conservation to the most relevant habitat types. There was a general decline of 40% in Northern European peatland bird population sizes in 1981-2014 (speed of decline 1.5%/year) largely driven by Finland, where populations declined almost 50% (2.0% annual decline). In Sweden and Norway, peatland bird populations declined by 20% during 1997-2014 (1.0% annual decline). In contrast, southern populations in Estonia and Latvia, where the majority of open peatlands are protected, showed a 40% increase during 1981-2014 (1.0% annual increase). The most important habitat characteristics preferred by common peatland species in Finland were openness and low tree height, while wetness proved to be an important feature for waders. Drainage of peatlands had clear negative effects on the densities of many species, with the only exception of rustic bunting, which specializes on edge habitats. Our findings call for more effective conservation actions in Northern European peatland habitats, especially in Finland where peatland drainage represents a major threat to biodiversity.

Forests used for timber production provide essential ecosystem services to society, as well as potential breeding habitat for bird communities. In southern Sweden, 90% of productive forest land is used for timber production and stands dominated by Norway spruce (Picea abies) constitute approximately 40% of the forested area. Due to their homogeneous structure, these spruce production forests are often regarded as depauperate. Despite this perception, knowledge about the biodiversity found in these stands is scarce. Here we synthesize the results of four separate bird surveys conducted within 35 spruce production stands of southern Sweden. The results are compared to recent population trends within the general study area. In total 49 bird species were recorded, with a strong difference in species composition between newly planted clear-cuts (forest age

Changes in forestry practices and on-going climate change may both have large impacts on forest bird populations. However, large-scale analyses of the effects of temporal changes in forest structure on forest bird numbers are largely lacking. We compared temporal trends from two Swedish nationwide long-term monitoring schemes, the Swedish Bird Survey (1998–2015) and the Swedish National Forestry Inventory (1983–2014), giving representative values for both forest and bird changes over an area of 35 million ha. Since 1998 the total area of middle-aged and mature forest increased by 6.4%. In parallel, several forest structures potentially beneficial to birds (dead wood, retention trees on clear cuts, multi-layer forests, old forest and broadleaved forest) increased somewhat in abundance, most likely as a result of legislation changes and increasing areas under forest certification schemes. Summer temperatures also increased, with warm summers dominating since 2002. In 1998–2015, the population sizes of 58 forest bird species on average increased, as did the number of species observed per route, with no general difference between forest specialists (16 species) and generalists (42 species). However, from around 2005, the positive trends in bird numbers and many forest structures have levelled out. An analysis of species population trends in relation to a measure of climate sensitivity (Species Temperature Index, STI) suggested that forest birds, just like Swedish birds in general, have indeed been affected by a warming climate. But given their STI, forest birds on average had more positive trends than non-forest birds, suggesting that other factors than climate have affected them positively. Strong candidate factors are the documented changes in forest quality and quantity. Whereas our data and analyses are correlational, and no firm conclusions on causality therefore can be drawn, it is reasonable to assume that the recent increases in forest quantity, forest quality, and summer temperatures, all have contributed to the general increase in forest bird numbers in Sweden. But the relative contribution of these driving forces remains to be determined. When it comes to the potentially positive effects of improving forest quality in terms of increases in old forest, stratification, retention trees and dead wood, it is noteworthy that many of the positive trends in forest structures since the mid-1990s seem to have ceased recently.

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.

Wind tunnels, in which birds fly against an artificially generated air flow, have since long been used to evaluate aerodynamic properties of steady bird flight. A new generation of wind tunnels has also allowed the many processes associated with migratory flights to be studied in captivity. We review how wind tunnel studies of aerodynamics and migratory performance together have helped advancing our understanding of bird migration. Current migration theory is based on the power-speed relationship of flight as well as flight range equations, both of which can be evaluated using birds flying in wind tunnels. In addition, and depending on wind tunnel properties, performance during gliding and climbing flight, and effects of air pressure, humidity and turbulence on bird flight has been measured. Long-distance migrant species have been flown repeatedly for up to 16 h non-stop, allowing detailed studies of the energy expenditure, fuel composition, protein turnover, water balance, immunocompetence and stress associated with sustained migratory flights. In addition, wind tunnels allow the fuelling periods between migratory flights to be studied from new angles. We end our review by suggesting several important topics for future wind tunnel studies, ranging from on of the key questions remaining, the efficiency at which chemical power in converted to mechanical power, to new useful avenues, such as improving and calibrating the techniques used for tracking of individual birds in the wild. (Less)

Aim Changes in climate and land use practices have been found to affect animal populations in different parts of the world. These studies have typically been conducted during the breeding season, whereas the non-breeding season (hereafter 'winter') has received much less attention. Changes in regional winter abundances could be caused by changes in overall population sizes and/or redistribution of populations. We tested these mechanisms for terrestrial winter bird population changes in Northern Europe and explored the role of climate change and species habitat preference.Location The Netherlands, Denmark, Sweden, Finland.Methods We used winter bird counts from four countries conducted annually between 15 December and 20 January in 1980/1981-2013/2014. We report national population trends for 50 species for which a trend could be calculated in at least three of the countries. We analysed country-specific population growth rates in relation to species' climatic summer and winter niches, habitat preference and migratory behaviour.Results Species breeding in colder (typically northern) areas showed more negative winter population trends than species breeding in warmer areas. Regional winter population trends were negatively correlated with characteristics of their winter climatic niche: populations in the colder part of their winter distribution increased in abundance, whereas populations in the warmer part of their winter distribution decreased. Woodland species tended to do better than farmland species. Migratory behaviour did not explain variation in population trends.Main conclusions The generally decreasing winter population trends of cold-dwelling breeding species probably reflect the general decline in population sizes of these species. In contrast, increasing winter population trends for populations in the colder parts of the winter distribution indicate a redistribution of wintering individuals towards the north-east. Both these patterns are likely caused by climate change.

The intensification of agriculture has resulted in more homogeneous landscapes and declines of many species associated with farmland or other semi-natural open habitats. In parallel, forestry has also intensified causing declines in many species associated with old-growth forests. While intensive forestry negatively affects forest species, it inadvertently creates new habitats such as clear-cuts, which attracts some farmland species. To understand the potential of clear-cuts as alternative habitat for farmland species, we need to know what makes clear-cuts attractive and whether they are suitable for reproduction and survival. We reviewed literature on the occurrence of farmland birds and butterflies in forest clear-cuts and synthesise the current knowledge on factors and characteristics affecting their occurrence. Many farmland birds and butterflies do indeed use clear-cuts, and have been found in clear-cuts up to ten years after felling. Clear-cut characteristics of importance include age, size, retention structures, land-use history and landscape composition. However, direct measures of resource abundance such as food and hostplants are often lacking. In addition to the potential benefit of individual clear-cuts, the total clear-cut area in forested regions is often large. Together with the fact that clear-cuts may be occupied by farmland species for several years, the potential of clear-cuts as alternative habitat for farmland biodiversity is substantial. Clear-cuts with a history as meadows, the presence of species of conservation importance, or shorter distance to farmland could for example be motivations for focusing conservation efforts on farmland species instead of forest species. Gaining more knowledge on how farmland species use clear-cuts, and what characteristics they depend on, could help inform management guidelines. We are no advocates for forest clear-cuts, but given their ubiquity in forested landscapes, the potential of clear-cuts as alternative habitats for species suffering from loss of suitable farmland habitats is worth serious attention from a conservation perspective.

Climate change is a major global threat to biodiversity with widespread impacts on ecological communities. Evidence for beneficial impacts on populations is perceived to be stronger and more plentiful than that for negative impacts, but few studies have investigated this apparent disparity, or how ecological factors affect population responses to climatic change. We examined the strength of the relationship between species-specific regional population changes and climate suitability trends (CST), using 30-year datasets of population change for 525 breeding bird species in Europe and the USA. These data indicate a consistent positive relationship between population trend and CST across the two continents. Importantly, we found no evidence that this positive relationship differs between species expected to be negatively and positively impacted across the entire taxonomic group, suggesting that climate change is causing equally strong, quantifiable population increases and declines. Species’ responses to changing climatic suitability varied with ecological traits, however, particularly breeding habitat preference and body mass. Species associated with inland wetlands responded most strongly and consistently to recent climatic change. In Europe, smaller species also appeared to respond more strongly, whilst the relationship with body mass was less clear-cut for North American birds. Overall, our results identify the role of certain traits in modulating responses to climate change and emphasise the importance of long-term data on abundance for detecting large-scale species’ responses to environmental changes.

Aims: Climate change is known to drive both the reshuffling of whole assemblages and range shifts of individual species. Less is known about how local colonizations and extinctions of individual species contribute to changes at the community level. Our aim was to estimate the contribution of individual species to a change in community composition attributed to climate change and to relate these species-specific contributions to species' commonness, climatic niche characteristics and life history traits most likely to influence species sensitivity to climate change. Location: Sweden. Methods: Focussing on birds, we analysed changes from 1998 to 2012 in the Community Temperature Index (CTI), a measure of the average climatic niche of a community. Using a jackknife approach we assessed the contribution of individual species to the temporal trend in CTI in four different regions across Sweden, controlling for habitat distribution. We further tested whether species contribution was related to population trends and rarity to identify species most vulnerable to climate change. Results: Community Temperature Index had increased over time with the greatest gains occurring in the north of the country, reflecting the larger temperature increases in this area. Changes in the regional CTI were driven both by warm-dwelling species colonizing new sites and by extirpations of cold-dwelling species. Furthermore, the community changes were influenced by both rare and common species. At the same time, the distribution changes of a large number of species were seemingly unaffected by climate change. Main conclusions: Both range expansion and contractions contributed to the relative increase of warm-dwelling species in Swedish bird communities. We successfully identified the climatic impacts on some of Sweden's rarest species, including cold-dwelling species in the mountainous north. Our approach may be an efficient tool to use when characterizing the impacts of climate change on species and communities.

Large-scale and population-wide monitoring of waders on their boreal and arctic breeding grounds has hitherto been lacking, mainly because logistics are truly challenging in regions with few ornithologists, vast areas and few roads. In Norway, Sweden and Finland (here ‘Fennoscandia’) there are now national monitoring schemes in place, aimed at tracking all bird species, which allows trends to be estimated for northern wader populations. We present joint Fennoscandian population trends for 24 wader species, covering the period 2002–2013 (in some cases somewhat shorter time periods). The data stem from 1263 routes in Norway, Finland, and the northern two thirds of Sweden, all situated north of 58°N. This area of one million km2 largely coincides with the boreal and arctic parts of Fennoscandia. The trends found are rather evenly distributed between strong increases and strong declines. Trends do not differ between short- and long-distance migrants, nor do they vary in relation to breeding latitude. Ringed Plover Charadrius hiaticula, Wood Sandpiper Tringa glareola, Green Sandpiper T. ochropus and Common Redshank T. totanus had significant positive trends, whereas Common Snipe Gallinago gallinago, Ruff Philomachus pugnax and Spotted Redshank Tringa erythropus were declining significantly in numbers. Trends could be calculated even for relatively uncommon breeding birds such as Temminck's Stint Calidris temminckii, Eurasian Dotterel Charadrius morinellus, Broad-billed Sandpiper Limicola falcinellus, Jack Snipe Lymnocryptes minimus and Red-necked Phalarope Phalaropus lobatus, although these trends build on few routes and individuals, and thus have low precision. The monitoring schemes in Norway and Finland are expected to increase in coverage in the coming years, with up to a total of 1555 northern routes being tracked when the schemes are fully developed. This should enable still more robust trend estimates for northern waders on their Fennoscandian breeding grounds to be calculated in the future.

Aim: To quantify avian distribution shifts and the extent of niche tracking in response to changing temperature and precipitation patterns. Location: Sweden. Methods: We used abundance monitoring data to quantify changes in bird species distributions between two time periods, 2000-02 and 2010-12. First we examined shifts at the level of whole distributions using population centroids in temperature, rainfall, altitude, latitude and longitude. We then characterized shifts in temperature and latitude at different parts of species ranges using species response curves (SRC). We accounted for yearly turnover in abundance and sampling effort, and compared the observed shifts with those expected under perfect niche tracking. Results: Most species demonstrated changes in their distributions over the last decade but not all were in response to weather. The degree to which species tracked their climatic niches and the dynamics driving these shifts varied considerably. Only 20% of species shifted in the direction expected given the temperature changes, while few showed a strong response to rainfall. Most shifts did not fully compensate for changes in temperature. Range changes were most evident at the leading edges, but there was some evidence for retractions of trailing edges. Amongst species that tracked temperature, those with southerly distributions were less successful at tracking changes than those in the north. Main conclusions: Swedish birds demonstrated highly dynamic distributions, with many rapid directional shifts occurring over the last decade. However, only a few species kept pace with observed climatic change. Species that did not track their climatic niche may be tolerant to ongoing climatic change or constrained by strong habitat requirements. We demonstrate that measuring range shifts along both environmental and geographic gradients can help disentangle drivers of distribution changes.

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.

Great Snipes Gallinago media spend about eight months per year in sub-Saharan Africa, but most aspects of their non-breeding ecology are poorly known. We analysed the seasonal pattern of appearance, flight feather moult (primaries and secondaries), and body mass in relation to age and sex, based on 3,247 birds collected by hunters in 1990–2000 in Benin, Gabon, Cameroon and the Democratic Republic of Congo. Adult males arrived in Africa from mid-August with a suspended flight feather moult. Adult females on average arrived somewhat later, and were about one month behind in the progress of flight feather moult. The adults of both sexes resumed moult immediately upon arrival. Flight feather moult was generally completed by the end of November in males, and end of December in females. Juvenile Great Snipes arrived later than adults and did not moult their flight feathers. The temporal pattern of occurrence at the different study sites suggests a general relocation of snipes from West Africa to Central Africa in October-December. Body masses did not differ between age groups and were generally low from August to February (155–170 g in males and 165–185 g in females). Males apparently departed northwards from late March to late April, and the females about two weeks later. Prior to departure, body mass increased on average with about 50 g (in both sexes), suggesting a departure fuel load of 31–33% above lean body mass. In periods of high and stable mass increase, fuel was deposited at rates of about 0.8–1.3 g/d (or 0.5–0.8% of lean body mass/d). The heaviest males and females had a total body mass of 242 and 250 g, respectively. This kind of traditional natural history data forms an important complement to the new type of data emerging from modern tracking techniques.

Large-scale multi-species data on population changes of alpine or arctic species are largely lacking. At the same time, climate change has been argued to cause poleward and uphill range shifts and the concomitant predicted loss of habitat may have drastic eff ects on alpine and arctic species. Here we present a multi-national bird indicator for the Fennoscandian mountain range in northern Europe (Finland, Sweden and Norway), based on 14 common species of montane tundra and subalpine birch forest. Th e data were collected at 262 alpine survey plots, mainly as a part of geographically representative national breeding bird monitoring schemes. Th e area sampled covers around 1/4 million km 2 , spanning 10 degrees of latitude and 1600 km in a northeast – southwest direction. During 2002 – 2012, nine of the 14 bird species declined signifi cantly in numbers, in parallel to higher summer temperatures and precipitation during this period compared to the preceding 40 yr. Th e population trends were largely parallel in the three countries and similar among montane tundra and subalpine birch forest species. Long-distance migrants declined less on average than residents and short-distance migrants. Some potential causes of the current decline of alpine birds are discussed, but since montane bird population sizes may show strong natural annual variation due to several factors, longer time series are needed to verify the observed population trends. Th e present Fennoscandian monitoring systems, which from 2010 onwards include more than 400 montane survey plots, have the capacity to deliver a robust bird indicator in the climate-sensitive mountainous regions of northernmost Europe for conservation purposes.

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.

Capsule Swedish breeding bird species that are predicted to experience a more suitable climate by 2050 are currently increasing in numbers. Aims To test, for a large set of breeding birds, if recent population trends mirror the changes in predicted climatic suitability across the Swedish distribution range, and to test if considering natal dispersal improves the fit of the predictive models. Methods Recent decadal population trends of 131 species of birds breeding in Sweden were compared to forecasted change in their future national distribution range by 2050, as simulated by suitability distribution models using climate and natal dispersal scenarios. Two other potential predictors of recent demographic trends are controlled for – namely habitat specialization and generation time. Results The 1998–2009 population trends were positively correlated with predicted changes in distribution range. Thus, forecasted changes in climatic suitability predict recent population trends. Accounting for the species-speci...

Maximizing the area under biodiversity‐related conservation measures is a main target of the European Union (EU) Biodiversity Strategy to 2020. We analyzed whether agrienvironmental schemes (AES) within EU common agricultural policy, special protected areas for birds (SPAs), and Annex I designation within EU Birds Directive had an effect on bird population changes using monitoring data from 39 farmland bird species from 1981 to 2012 at EU scale. Populations of resident and short‐distance migrants were larger with increasing SPAs and AES coverage, while Annex I species had higher population growth rates with increasing SPAs, indicating that SPAs may contribute to the protection of mainly target species and species spending most of their life cycle in the EU. Because farmland birds are in decline and the negative relationship of agricultural intensification with their population growth rates was evident during the implementation of AES and SPAs, EU policies seem to generally attenuate the declines of farmland bird populations, but not to reverse them. PECBMS is supported financially by the RSPB and the European Commission. TT was supported by Institutional Research Plan (RVO: 68081766), SH and LB were supported by EUBON project (308454; FP7‐ENV‐2012 European Commission) and the TRUSTEE project (RURAGRI ERA‐NET 235175), and AL received financial support from the Academy of Finland (project 275606).

Farmland birds in Europe have declined in numbers from the last decades, as reflected in the Farmland Bird Index (FBI), a multi-species indicator of population trends adopted by the European Union. Some of these farmland bird species breed to a varying degree also in non-farmland habitats: however, the indicator is calculated using trends estimated from all birds counted of each species, irrespective of habitat. Consequently, when interpreting indicators of farmland birds, it is important to know the influence of including the trends of the populations in non-farmland habitats. We estimated national and regional bird-habitat relationships and population trends of 14 of the species included in the Swedish version of FBI. Bird data came from the Swedish Bird Survey (1996-2008) including 716 separate routes, and detailed habitat data came from the Swedish Board of Agriculture (1999-2008). The proportion of individuals that occurred in non-farmland habitats ranged from 2% to 96%. Local densities were generally highest in farmland habitats, but because only 7-8% of the surface of Sweden is farmland, even low local densities in non-farmland sum up to high national totals. With few exceptions, the temporal trends of species were independent of the amount of farmland on routes, suggesting similar trends within and outside farmland. Accordingly, incorporating data on farmland birds from non-farmland habitats had only a limited effect on the FBI. But given that habitat-specific trends may well occur, it is important that multi-species indices are checked for potential habitat-specific differences among the included species. (C) 2012 Elsevier Ltd. All rights reserved.