Specialization is a concept based on a broad theoretical framework developed by evolutionary biologists and ecologists. In the past 10 years, numerous studies have reported that – in many contexts – generalist species are “replacing” specialist species. We review recent research on the concept of the ecological niche and species specialization, and conclude that (1) the observed worldwide decline in specialist species is predicted by niche theory, (2) specialist declines cause “functional homogenization” of biodiversity, and (3) such homogenization may be used to measure the impact of disturbance on communities. Homogenization at the community level could alter ecosystem functioning and productivity, as well as result in the deterioration of ecosystem goods and services. We propose community‐level specialization as an indicator of the impact of global changes (habitat and climate disturbances) on biodiversity.

Climate changes have profound effects on the distribution of numerous plant and animal species 1-3 . However, whether and how different taxonomic groups are able to track climate changes at large spatial scales is still unclear. Here, we measure and compare the climatic debt accumulated by bird and butterfly communities at a European scale over two decades (1990)(1991)(1992)(1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008). We quantified the yearly change in community composition in response to climate change for 9,490 bird and 2,130 butterfly communities distributed across Europe 4 . We show that changes in community composition are rapid but different between birds and butterflies and equivalent to a 37 and 114 km northward shift in bird and butterfly communities, respectively. We further found that, during the same period, the northward shift in temperature in Europe was even faster, so that the climatic debts of birds and butterflies correspond to a 212 and 135 km lag behind climate. Our results indicate both that birds and butterflies do not keep up with temperature increase and the accumulation of different climatic debts for these groups at national and continental scales.Species are not equally at risk when facing climate change. Several species-specific attributes have been identified as increasing species' vulnerability to climate change, including diets, migratory strategy, main habitat types and ecological specialization [5][6][7] . Moreover, although phenotypic plasticity may enable some species to respond rapidly and effectively to climate change 8,9 , others may suffer from the induced spatial mismatch and temporal mistiming with their resources 10,11 . For instance, species such as great tits and flycatchers have been shown to become desynchronized with their main food supply during the nesting season 12 .However, beyond individual species' fates, climate change should also affect species interactions and the structure of species assemblages within and across different taxonomic groups over large spatial scales [13][14][15] . For instance, ectotherms should be more directly affected by climate warming and taxonomic groups with short generation time should favour faster evolutionary responses to selective pressures induced by climate changes 13 . Yet, whether different taxonomic groups are tracking climate change at the same rate over large areas is still unclear, and methods to routinely assess the mismatch between temperature increases and biodiversity responses at different spatial scales are still missing 16 .Here, we used extensive monitoring data of birds and butterflies distributed across Europe to assess whether, regardless of their species-specific characteristics, organisms belonging to a given group are responding more quickly or more slowly than organisms belonging to another group over large areas. We characterized bird and butterfly communities in 9,490 and 2,130 sample sites respectively by their community temperature index (CTI) for ea...

Summary1. Ecological specialization is one of the main concepts in ecology and conservation. However, this concept has become highly context-dependent and is now obscured by the great variability of existing definitions and methods used to characterize ecological specialization. 2. In this study, we clarify this concept by reviewing the strengths and limitations of different approaches commonly used to define and measure ecological specialization. We first show that ecological specialization can either be considered as reflecting species' requirements or species' impacts. We then explain how specialization depends on species-specific characteristics and on local and contingent environmental constraints. We further show why and how ecological specialization should be scaled across spatial and temporal scales, and from individuals to communities. 3. We then illustrate how this review can be used as a practical toolbox to classify widely used metrics of ecological specialization in applied ecology, depending on the question being addressed, the method used, and the data available. 4. Synthesis and applications. Clarifying ecological specialization is useful to make explicit connections between several fields of ecology using the niche concept. Defining this concept and its practical metrics is also a crucial step to better formulate predictions of scientific interest in ecology and conservation. Finally, understanding the different facets of ecological specialization should facilitate to investigate the causes and consequences of biotic homogenization and to derive relevant indicators of biodiversity responses to land-use changes.

Range shifts of many species are now documented as a response to global warming. But whether these observed changes are occurring fast enough remains uncertain and hardly quantifiable. Here, we developed a simple framework to measure change in community composition in response to climate warming. This framework is based on a community temperature index (CTI ) that directly reflects, for a given species assemblage, the balance between low-and high-temperature dwelling species. Using data from the French breeding bird survey, we first found a strong increase in CTI over the last two decades revealing that birds are rapidly tracking climate warming. This increase corresponds to a 91 km northward shift in bird community composition, which is much higher than previous estimates based on changes in species range edges. During the same period, temperature increase corresponds to a 273 km northward shift in temperature. Change in community composition was thus insufficient to keep up with temperature increase: birds are lagging approximately 182 km behind climate warming. Our method is applicable to any taxa with large-scale survey data, using either abundance or occurrence data. This approach can be further used to test whether different delays are found across groups or in different land-use contexts.

Each species generally has a close relationship with one or more habitats and can therefore be classified as either specialist or generalist. We studied whether specialist and generalist species are spatially distributed independently of each other. Repeating the analysis for 100 of the most frequent terrestrial bird species recorded over the 10 000 sampled sites of the French Breeding Bird survey, we found that specialists were more abundant if the rest of the community was specialized, and that the inverse was also true. This pattern was far subtler than just a simple dichotomy: most species actually presented a maximum abundance at a value of community specialization similar to their own level of specialization. Bird communities appear very well defined along a specialist-generalist gradient. We believe this pattern becomes more apparent with habitat degradation. The consequences on both ecological services and community resilience may well be considerable.

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Aim Worldwide, functional homogenization is now considered to be one of the most prominent forms of biotic impoverishment induced by current global changes. Yet this process has hardly been quantified on a large scale through simple indices, and the connection between landscape disturbance and functional homogenization has hardly been established. Here we test whether changes in land use and landscape fragmentation are associated with functional homogenization of bird communities at a national scale. Location France.Methods We estimated functional homogenization of a community as the average specialization of the species present in that community. We studied the spatial variation of this community specialization index (CSI) using 1028 replicates from the French Breeding Bird Survey along spatial gradients of landscape fragmentation and recent landscape disturbance, measured independently, and accounting for spatial autocorrelation. ResultsThe CSI was very sensitive to both measures of environmental degradation: on average, 23% of the difference in the CSI values between two sample sites was attributed to the difference in fragmentation and the disturbance between sites. This negative correlation between CSI and sources of landscape degradation was consistent over various habitats and biogeographical zones. Main conclusionsWe demonstrate that the functional homogenization of bird communities is strongly positively correlated to landscape disturbance and fragmentation. We suggest that the CSI is particularly effective for measuring functional homogenization on both local and global scales for any sort of organism and with abundance or presence-absence data.

Few studies have examined how life history traits and the climate envelope influence the ability of species to respond to climate change and habitat degradation. In this study, we test whether 18 species-specific variables, related to the climate envelope, ecological envelope and life history, could predict recent population trends (over 17 years) of 71 common breeding bird species in France. Habitat specialists were declining at a much higher rate than generalists, a sign that habitat quality is decreasing globally. The lower the thermal maximum (temperature at the hot edge of the climate envelope), the more negative are the population trends and the less tolerant these species are climate warming, regardless of the thermal range over which these species occur. The life history trait 'the number of broods per year' was positively related to recent trends, suggesting that single-brooded species might be more sensitive to advances in food peak due to climate change, as it increases the risk of mistiming their single-breeding event. Annual fecundity explained long-term declines, as it is a good proxy for most other demographic rates, with shorter-lived species being more sensitive to global change: individuals of species with higher fecundity might have too short a life to learn to adapt to directional changes in their environment. Finally, there was evidence that natal dispersal was a predictor of recent trends, with species with high natal dispersal experiencing smaller population declines than species with low natal dispersal. This is expected if the higher the natal dispersal, the larger the ability to shift spatially when facing changes in local habitat or climate, in order to track optimal conditions and adapt to global change. Identifying decline-promoting factors allow us to infer mechanisms responsible for observed declines in wild bird populations facing global change, and by doing so allow for a more pre-emptive approach to conservation planning.