Maximizing regional biodiversity requires a mosaic of protection levels.

Protected areas are the flagship management tools to secure biodiversity from anthropogenic impacts. However, the extent to which adjacent areas with distinct protection levels host different species numbers and compositions remains uncertain. Here, using reef fishes, European alpine plants, and North American birds, we show that the composition of species in adjacent Strictly Protected, Restricted, and Non-Protected areas is highly dissimilar, whereas the number of species is similar, after controlling for environmental conditions, sample size, and rarity. We find that between 12% and 15% of species are only recorded in Non-Protected areas, suggesting that a non-negligible part of regional biodiversity occurs where human activities are less regulated. For imperiled species, the proportion only recorded in Strictly Protected areas reaches 58% for fishes, 11% for birds, and 7% for plants, highlighting the fundamental and unique role of protected areas and their environmental conditions in biodiversity conservation.

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.

Short-term climate-induced change in French plant communities.

Latitudinal and altitudinal range shifts in response to climate change have been reported for numerous animal species, especially those with high dispersal capacities. In plants, the impact of climate change on species distribution or community composition has been documented mainly over long periods (decades) and in specific habitats, often forests. Here, we broaden the results of such long-term, focused studies by examining climate-driven changes in plant community composition over a large area (France) encompassing multiple habitat types and over a short period (2009-2017). To this end, we measured mean community thermal preference, calculated as the community-weighted mean of the Ellenberg temperature indicator value, using data from a standardized participatory monitoring scheme. We report a rapid increase in the mean thermal preference of plant communities at national and regional scales, which we relate to climate change. This reshuffling of plant community composition corresponds to a relative increase in the abundance of warm- versus cold-adapted species. However, support for this trend was weaker when considering only the common species, including common annuals. Our results thus suggest for the first time that the response of plant communities to climate change involves subtle changes affecting all species rare and common, which can nonetheless be detected over short time periods. Whether such changes are sufficient to cope with the current climate warming remains to be ascertained.

Where do they go? The effects of topography and habitat diversity on reducing climatic debt in birds.

The spatial tracking of climatic shifts is frequently reported as a biodiversity response to climatic change. However, species' range shifts are often idiosyncratic and inconsistent with climatic shift predictions. At the community scale, this discrepancy can be measured by comparing the spatial shift in the relative composition of cold- vs. warm-adapted species in a local assemblage [the community temperature index (CTI)] with the spatial shift in temperature isotherms. While the local distribution of climate change velocity is a promising approach to downscaling climate change pressure and responses, CTI velocity has only been investigated on a continental or national scale. In this study, we coupled French Breeding Bird Survey data, collected from 2133 sites monitored between 2001 and 2012, with climatic data in order to estimate the local magnitude and direction of breeding season temperature shift, CTI shift, and their spatiotemporal divergence - the local climatic debt. We also tested whether landscape characteristics that are known to affect climate velocity and spatial tracking of climate change mediated the climatic debt on the local scale. We found a clear spatial structure, together with heterogeneity in both temperature and CTI spatial shifts. Local climatic debt decreased as the elevation, habitat diversity, and the naturalness of the landscape increased. These results suggest the complementary effects of the local topographic patterns sheltering more diverse microclimates and the increasing permeability of natural and diversified landscape. Our findings suggest that a more nuanced evaluation of spatial variability in climatic and biotic shifts is necessary in order to properly describe biodiversity responses to climate change rather than the oversimplified descriptions of uniform poleward shifts.

Rapid adjustment of bird community compositions to local climatic variations and its functional consequences.

The local spatial congruence between climate changes and community changes has rarely been studied over large areas. We proposed one of the first comprehensive frameworks tracking local changes in community composition related to climate changes. First, we investigated whether and how 12 years of changes in the local composition of bird communities were related to local climate variations. Then, we tested the consequences of this climate-induced adjustment of communities on Grinnellian (habitat-related) and Eltonian (function-related) homogenization. A standardized protocol monitoring spatial and temporal trends of birds over France from 2001 to 2012 was used. For each plot and each year, we used the spring temperature and the spring precipitations and calculated three indices reflecting the thermal niche, the habitat specialization, and the functional originality of the species within a community. We then used a moving-window approach to estimate the spatial distribution of the temporal trends in each of these indices and their congruency with local climatic variations. Temperature fluctuations and community dynamics were found to be highly variable in space, but their variations were finely congruent. More interestingly, the community adjustment to temperature variations was nonmonotonous. Instead, unexplained fluctuations in community composition were observed up to a certain threshold of climate change intensity, above which a change in community composition was observed. This shift corresponded to a significant decrease in the relative abundance of habitat specialists and functionally original species within communities, regardless of the direction of temperature change. The investigation of variations in climate and community responses appears to be a central step toward a better understanding of climate change effects on biodiversity. Our results suggest a fine-scale and short-term adjustment of community composition to temperature changes. Moreover, significant temperature variations seem to be responsible for both the Grinnellian and Eltonian aspects of functional homogenization.

Asynchrony of taxonomic, functional and phylogenetic diversity in birds.

AIM: We assessed the temporal trends of taxonomic, functional and phylogenetic diversities in the French avifauna over the last two decades. Additionally, we investigated whether and how this multifaceted approach to biodiversity dynamics can reveal an increasing similarity of local assemblages in terms of species, traits and/or lineages. LOCATION: France. METHODS: We analysed a large-scale dataset that recorded annual changes in the abundance of 116 breeding birds in France between 1989 and 2012. We decomposed and analysed the spatio-temporal dynamics of taxonomic, phylogenetic and functional diversities and each of their α-, ß- and γ-components. We also calculated the trend in the mean specialization of bird communities to track the relative success of specialist versus generalist species within communities during the same period. RESULTS: We found large variation within and among the temporal trends of each biodiversity facet. On average, we found a marked increase in species and phylogenetic diversity over the period considered, but no particular trend was found for functional diversity. Conversely, changes in ß-diversities for the three facets were characterized by independent and nonlinear trends. We also found a general increase in the local occurrence and abundance of generalist species within local communities. MAIN CONCLUSIONS: These results highlight a relative asynchrony of the different biodiversity facets occurring at large spatial scales. We show why a multifaceted approach to biodiversity dynamics is needed to better describe and understand changes in community composition in macroecology and conservation biogeography.

Multifaceted diversity-area relationships reveal global hotspots of mammalian species, trait and lineage diversity.

AIM: To define biome-scale hotspots of phylogenetic and functional mammalian biodiversity (PD and FD, respectively) and compare them to 'classical' hotspots based on species richness (SR) only. LOCATION: Global. METHODS: SR, PD & FD were computed for 782 terrestrial ecoregions using distribution ranges of 4616 mammalian species. We used a set of comprehensive diversity indices unified by a recent framework that incorporates the species relative coverage in each ecoregion. We build large-scale multifaceted diversity-area relationships to rank ecoregions according to their levels of biodiversity while accounting for the effect of area on each diversity facet. Finally we defined hotspots as the top-ranked ecoregions. RESULTS: While ignoring species relative coverage led to a relative good congruence between biome top ranked SR, PD and FD hotspots, ecoregions harboring a rich and abundantly represented evolutionary history and functional diversity did not match with top ranked ecoregions defined by species richness. More importantly PD and FD hotspots showed important spatial mismatches. We also found that FD and PD generally reached their maximum values faster than species richness as a function of area. MAIN CONCLUSIONS: The fact that PD/FD reach faster their maximal value than SR may suggest that the two former facets might be less vulnerable to habitat loss than the latter. While this point is expected, it is the first time that it is quantified at global scale and should have important consequences in conservation. Incorporating species relative coverage into the delineation of multifaceted hotspots of diversity lead to weak congruence between SR, PD and FD hotspots. This means that maximizing species number may fail at preserving those nodes (in the phylogenetic or functional tree) that are relatively abundant in the ecoregion. As a consequence it may be of prime importance to adopt a multifaceted biodiversity perspective to inform conservation strategies at global scale.

[Environmental protection: ethical and scientific sources of tension]. / La protection de la nature: une double tension éthique et scientifique.

Environmental ethic is complex, dynamic, and related to cultural contexts that are neither given once and for all, nor valid for all people. Yet, the current biodiversity crisis tends to be followed by the spread of universal values related to nature and its protection. Far from standardizing the pluralism of these values, this universal trend leads to two major tensions in the field of nature conservation. The first concerns the scientific or epistemic values: nature or "biodiversity" is altogether being reduced in sub-categories or by contrast considered as a complex object by modern ecological science. Biodiversity is indeed both increasingly considered in terms of its complexity and unpredictability, or on the contrary, as a collection of natural objects that can and must be managed with easily quantifiable indicators. The second tension concerns ethical values. These values tend to be reduced and simplified in equating nature with "services" that can benefit human societies. But this utilitarian reduction also hides a diversification of ethical values. This article aims to analyze what characterizes this contemporary ethical and scientific floating with respect to the values involved in the protection of nature.

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.

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.

Using phylogenies in conservation: new perspectives.

The 2011 meeting of the European Ecological Federation took place in Ávila, Spain, from 26th September to 29th September. The French Ecological Society (SFE) and the Foundation for Research on Biodiversity (FRB) sponsored a session entitled 'Evolutionary history, ecosystem function and conservation biology: new perspectives'. We report on the main insights obtained from this symposium.

Titmice are a better indicator of bird density in Northern European than in Western European forests.

Population sizes of many birds are declining alarmingly and methods for estimating fluctuations in species' abundances at a large spatial scale are needed. The possibility to derive indicators from the tendency of specific species to co-occur with others has been overlooked. Here, we tested whether the abundance of resident titmice can act as a general ecological indicator of forest bird density in European forests. Titmice species are easily identifiable and have a wide distribution, which makes them potentially useful ecological indicators. Migratory birds often use information on the density of resident birds, such as titmice, as a cue for habitat selection. Thus, the density of residents may potentially affect community dynamics. We examined spatio-temporal variation in titmouse abundance and total bird abundance, each measured as biomass, by using long-term citizen science data on breeding forest birds in Finland and France. We analyzed the variation in observed forest bird density (excluding titmice) in relation to titmouse abundance. In Finland, forest bird density linearly increased with titmouse abundance. In France, forest bird density nonlinearly increased with titmouse abundance, the association weakening toward high titmouse abundance. We then analyzed whether the abundance (measured as biomass) of random species sets could predict forest bird density better than titmouse abundance. Random species sets outperformed titmice as an indicator of forest bird density only in 4.4% and 24.2% of the random draws, in Finland and France, respectively. Overall, the results suggest that titmice could act as an indicator of bird density in Northern European forest bird communities, encouraging the use of titmice observations by even less-experienced observers in citizen science monitoring of general forest bird density.

Waders in winter: long-term changes of migratory bird assemblages facing climate change.

Effects of climate change on species occupying distinct areas during their life cycle are still unclear. Moreover, although effects of climate change have widely been studied at the species level, less is known about community responses. Here, we test whether and how the composition of wader (Charadrii) assemblages, breeding in high latitude and wintering from Europe to Africa, is affected by climate change over 33 years. We calculated the temporal trend in the community temperature index (CTI), which measures the balance between cold and hot dwellers present in species assemblages. We found a steep increase in the CTI, which reflects a profound change in assemblage composition in response to recent climate change. This study provides, to our knowledge, the first evidence of a strong community response of migratory species to climate change in their wintering areas.

Spatial mismatch and congruence between taxonomic, phylogenetic and functional diversity: the need for integrative conservation strategies in a changing world.

Functional and phylogenetic diversity are increasingly quantified in various fields of ecology and conservation biology. The need to maintain diversity turnover among sites, so-called beta-diversity, has also been raised in theoretical and applied ecology. In this study, we propose the first comprehensive framework for the large-scale mapping of taxonomic, phylogenetic and functional diversity and of their respective turnover. Using high-resolution data on the spatial distribution and abundance of birds at a country scale, we disentangled areas of mismatches and congruencies between biodiversity components. We further revealed unequal representation of each component in protected areas: functional diversity was significantly under-represented whereas taxonomic diversity was significantly over-represented in protected areas. Our results challenge the use of any one diversity component as a surrogate for other components and stress the need to adopt an integrative approach to biodiversity conservation.

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.

On the relative importance of space and environment in farmland bird community assembly.

The relative contribution of ecological processes in shaping metacommunity dynamics in heavily managed landscapes is still unclear. Here we used two complementary approaches to disentangle the role of environment and spatial effect in farmland bird community assembly in an intensive agro-ecosystem. We hypothesized that the interaction between habitat patches and dispersal should play a major role in such unstable and unpredictable environments. First, we used a metacommunity patterns analysis to characterize species co-occurrences and identify the main drivers of community assembly; secondly, variation partitioning was used to disentangle environmental and geographical factors (such as dispersal limitation) on community structure and composition. We used high spatial resolution data on bird community structure and composition distributed among 260 plots in an agricultural landscape. Species were partitioned into functional classes, and point count stations were classified according to landscape characteristics before applying metacommunity and partitioning analyses within each. Overall we could explain around 20% of the variance in species composition in our system, revealing that stochasticity remains very important at this scale. However, this proportion varies depending on the scale of analysis, and reveals potentially important contributions of environmental filtering and dispersal. These conclusions are further reinforced when the analysis was deconstructed by bird functional classes or by landscape habitat classes, underlining trait-related filters, thus reinforcing the idea that wooded areas in these agroecosystems may represent important sources for a specific group of bird species. Our analysis shows that deconstructing the species assemblages into separate functional groups and types of landscapes, along with a combination of analysis strategies, can help in understanding the mechanisms driving community assembly.

Birds are tracking climate warming, but not fast enough.

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.