Sylvain Mollier

Georges Kunstler

Jean-Luc Dupouey

Laurent Bergès

Historical landscape matters for threatened species in French mountain forests

Ancient forests are known to host a biodiversity of high ecological distinctiveness and are likely to provide habitat for red-listed species. Yet, few studies have investigated the role of forest continuity for the conservation of threatened species. We used species-presence data on red-listed species from 12 taxonomic groups (Spermatophyta, Pteridophyta, Bryophyta, Lichens, Chiroptera, Aves, Squamata, Amphibia, Coleoptera, Lepidoptera, Odonata and Orthoptera) to ascertain if ancient forests are an important habitat for threatened species in five mountain and subalpine protected areas in France. We compared the effect of the amount of historical forest (1853-1860) with the effect of the amount of current forest on the distribution of red-listed species in six circular buffers landscape ranging in radius from 100 to 1500m. We showed that the amount of historical forest in the landscape had a positive effect on forest Spermatophyta, Bryophyta, Coleoptera and edge forest Pteridophyta with a better predictive power than current forest area, highlitghing a colonization credit in recent forests. Conversely, edge-forest lepidopterans were more negatively affected by historical than by current forest area, highlighting an extinction debt in recent forests. Our findings underline that implementing protective measures of ancient forests could be a better strategy than afforestation to preserve threatened forest species in mountain and subalpine forest landscapes.

I. Introduction

Forests with a long temporal continuity, i.e. forests with no change in land use for at least 150 years, also known as ancient forests (AF) as opposed to recent forests (RF), have particular conservation value because they host species guilds with low dispersal and competitive abilities that have trouble persisting after forest/habitat destruction [START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF][START_REF] Hermy | Legacies of the past in the present-day forest biodiversity: a review of past landuse effects on forest plant species composition and diversity[END_REF]. Even though the total surface area of temperate forests is currently expanding [START_REF] Keenan | Dynamics of global forest area: Results from the FAO Global Forest Resources Assessment 2015[END_REF][START_REF] Mather | The Forest Transition[END_REF]), AF area is still eroding. Indeed, it is estimated that 10 to 40% of the forests present in the 19 th century have disappeared in European countries today [START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF]. Despite this insidious erosion and behind the recommendations of the IUCN (IUCN 2016), only two European countries (Great Britain and Belgium) have adopted measures to protect AFs [START_REF] Goldberg | The ancient woodland concept as a practical conservation tool in Great Britain[END_REF][START_REF] Kervyn | La gestion patrimoniale des forêts anciennes de Wallonie (Belgique)[END_REF]. Species extinction probability is strongly related to population size, disturbances and the species' functional traits [START_REF] Fischer | Landscape modification and habitat fragmentation: a synthesis[END_REF][START_REF] Davies | The macroecology and macroevolution of plant species at risk[END_REF]. Therefore, extinction risk is likely to increase for species with low mobility, low dispersal and competition abilities and high habitat specialisation [START_REF] Kotiaho | Predicting the risk of extinction from shared ecological characteristics[END_REF][START_REF] Cooke | Ecological distinctiveness of birds and mammals at the global scale[END_REF]). In addition, rare species are particularly sensitive to past human activities [START_REF] Lavergne | Environmental and human factors influencing rare plant local occurrence, extinction and persistence: a 115-year study in the Mediterranean region[END_REF] and the proportion of extinct or threatened species is better explained by 1900's indicators of human activities than by current one, showing a time lag effect [START_REF] Gosselin | On the time lag between human activity and biodiversity in Europe at the national scale[END_REF]. Threatened species may therefore share many traits with ancient forest species. However, whereas many studies have highlighted the high ecological distinctiveness of AF biodiversity, only a few focused on the importance of these forests for threatened species [START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF]. Two studies found a positive correlation between ancient forest species richness and threatened species richness for bryophytes, lichens, macro fungi, beetles, butterflies and spiders, one in Czech Republic [START_REF] Hofmeister | Shared affinity of various forest-dwelling taxa point to the continuity of temperate forests[END_REF] and the other in Sweden [START_REF] Fritz | Does forest continuity matter in conservation? -A study of epiphytic lichens and bryophytes in beech forests of southern Sweden[END_REF]. However, these studies did not directly characterise forest continuity and only studied correlations between biodiversity indices. Only [START_REF] Flensted | Red-listed species and forest continuity -A multi-taxon approach to conservation in temperate forests[END_REF] actually evaluated the effect of past landscape context on the distribution of threatened forest species. The authors showed that threatened forest species richness in Danish forests was better correlated with historical forest area (mapped between 1760 and 1820) than with current forest area for mammals, saproxylic beetles, butterflies, vascular plants and four groups of fungi. However, this study was carried out in a highly fragmented landscape (14% of Denmark is currently forested) and did not explore the response of non-forest species, although they might react differently to the temporal continuity of the forest and be relevant for the implementation of conservation measures. Indeed, open-habitat dependent species can persist in recent forests for several reasons: (1) the canopy cover may remain rather low, (2) they are close to the forest edge [START_REF] Harper | Edge Influence on Forest Structure and Composition in Fragmented Landscapes[END_REF], and (3) their occurrence may be related to an extinction debt [START_REF] Jackson | Balancing biodiversity in a changing environment: extinction debt, immigration credit and species turnover[END_REF][START_REF] Milberg | Land-use history influence the vegetation in coniferous production forests in southern Sweden[END_REF].

To date, the role of past land use on forest biodiversity has rarely been studied in mountain forests [START_REF] Janssen | Influences relatives de l'ancienneté et de la maturité sur la biodiversité: implications pour la conservation en forêts de montagne[END_REF][START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF]. These forests have globally been less affected by deforestation and have undergone less intense sylvicultural management than their lowland counterparts, which might lead to weaker impacts on threatened species than those observed by [START_REF] Flensted | Red-listed species and forest continuity -A multi-taxon approach to conservation in temperate forests[END_REF].

Our aim was to analyse the response of threatened species from multiple taxonomic groups to historical and current landscape contexts, using species observations in forest only. We used a large dataset of threatened forest and non-forest species presence from 12 taxonomic groups (Spermatophyta, Pteridophyta, Bryophyta, lichens, Chiroptera, Aves, Squamata, Amphibia, Coleoptera, Lepidoptera, Odonata and Orthoptera) in five protected areas in the French mountains. We assessed the probability of observing a threatened species within each taxonomic group as a function of the amount of historical or current forest in concentric circular buffers of different radii. More specifically, we tested the following hypothesis: 1-Due to habitat limitation, the occurrence of threatened forest species should increase with the increasing amount of current forest in the landscape, while the occurrence of non-forest species should decrease. 2-A delayed response to land-use change would occur in recent forests (former open lands) due to dispersal and recruitment limitations of threatened forest species (colonisation credit) and persistence of threatened non-forest species (extinction debt). Thus, the occurrence of threatened forest species should increase with the amount of historical forest in the landscape, and should be better predicted by the amount of historical forest than by the amount of current forest. Conversely, the occurrence of non-forest species should increase with the amount of historical open land in the landscape (and thus should decrease with the amount of historical forest in the landscape), and should be better predicted by the amount of historical open land than by the amount of current open land.

II. Materials and methods

II.1. Study area

The study area encompassed five National Parks (NP) in the southern half of France. The Vanoise, the Ecrins and the Mercantour NPs cover a large part of the French Alps and benefit from a mountain climate with a continental influence. The Cevennes and the Pyrenees NPs cover part of the Massif Central and the Pyrenean range and have a mountain climate with, respectively, a mediterranean and oceanic influence. The study covered all the elevation range where forest is naturally present. Current forest cover in the five parks ranges from 19% to 68%, while historical forest cover ranges from 10 to 17% (Table 1). Deforestation represents 5.5% to 17% of the historical forested area but forest is currently in expansion and increased by +97% to +347% since 1853-1864. The Vanoise NP displayed the lowest forest cover with the largest share of AFs, while the Cevennes NP had the highest forest cover with the smallest share of AFs. Conifers dominated in the Vanoise, Ecrins and Mercantour NPs, while broadleaves dominated in the Cevennes and Pyrenees NPs.

II.2. Species presence data

We prepared our database by assembling existing databases from the National Park managers, the National Botanical Conservatories and the French Nature and Landscape Information System (https://openobs.mnhn.fr/). These data are based on opportunistic observations and do not contain information on species absence but they include a high number of observations of red-listed species, which are invaluable for describing the distribution of these species. Twelve taxonomic groups were considered: Spermatophyta, Pteridophyta, Bryophyta, lichens, Chiroptera, Aves, Squamata, Amphibia, Coleoptera, Lepidoptera, Odonata and Orthoptera. We selected observations made from January 2000 to July 2018 and retained only those made in closed forest (i.e. tree cover ≥ 40 % according to the French National Forest map BD FORET ® V2) at least 10 m from the forest edge and more than 1 500 m from the boundaries of the study area. Data were thinned using spThin R package [START_REF] Aiello-Lammens | spThin: an R package for spatial thinning of species occurrence records for use in ecological niche models[END_REF] by setting the minimum distance between observations of each species to 200m for birds and chiropterans and 100m for other taxonomic groups. Species classified as "Vulnerable", "Endangered" and "Critically Endangered" in the IUCN regional red lists were considered threatened. If the species status was not available at the regional level (no regional list, species classified "Data Deficient" or omitted from the regional list), we used the national or European red list status instead [START_REF] Iucn | IUCN Red List categories and criteria, version 3.1[END_REF]). This information was derived from [START_REF] Gargominy | Base de connaissance "Statuts" des espèces en France[END_REF] and synonyms were solved according to the official French taxonomic nomenclature [START_REF] Gargominy | TAXREF v14, référentiel taxonomique pour la France : méthodologie, mise en oeuvre et diffusion[END_REF]. As there is no official national or regional IUCN red list for lichens in France, we used the information provided by Roux (2020). Based on the references listed in Appendix S1, the species were classified into three categories (Schneider et al. 2021): forest-dependent species (forest species: "FS"), open habitatdependent species (non-forest species: "NFS") and forest-edge or generalist species (edge species: "ES").

As the dataset contained only presence data, we generated a set of pseudo-absences at the nodes of a 200 x 200 m grid. Then, we applied a list of filters to only keep: (1) pseudo-absences located in sectors (on a 2x2 km grid) with at least one observation of a threatened or non-threatened species from the corresponding taxonomic group, (2) pseudo-absences located in closed forests, and (3) pseudo-absences located more than 10 m from the forest edge and more than 1500 m from the boundaries of the study area.

II.3. Environmental factors

Large-scale context may determine local responses but the landscape scale of effect is difficult to determine a priori [START_REF] Avon | Landscape effects on plants in forests: Large-scale context determines local plant response[END_REF]. One common method is to measure landscape characteristics at different nested spatial scales and then determine which scale best explains the ecological response to the landscape context [START_REF] Jackson | Are ecologists conducting research at the optimal scale?[END_REF]. We therefore calculated the percentage of historical or current forest area in six circular buffers with radii of 100, 250, 500, 750, 1 000 and 1 500 m (noted B100 to B1500) for each threatened species observation point and each pseudo-absence point. We calculated the forested area in the current landscape using the BD FORET ® V2 map (consistent with the FAO's definition of "forest", FAO 2012), drawn in our study area between 2014 and 2018 by the French National Geographical Institute. We identified historical forests using the Ordnance survey maps drawn between 1853 and 1864, which are the reference in France for identifying long-continuity forests [START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF]. All the maps had previously been vectorised and georeferenced according to [START_REF] Favre | Digitalisation des cartes anciennes. Manuel pour la vectorisation de l'usage des sols et le géoréférencement des minutes 1:40 000 de la carte d'Etat-Major[END_REF] by the National Research Institute for Agriculture, food and the Environment (INRAE) and the National Parks [START_REF] Thomas | Long Term Changes in Forest Landscapes in Five National Parks in Metropolitan France and the Future Champagne and Bourgogne National Forest Park[END_REF]. We also included elevation (from the French DEM BD ALTI ® 25 m) and stand tree species composition (coniferous/deciduous/mixed, from the BD FORET ® V2 forest map) to control for local biotic and abiotic conditions. To control for potential sampling bias, we also included slope and distance to the closest path in the analyses.

The data were processed with the sf (Pebesma 2018) and raster (Hijmans 2020) packages from the R 4.1.1 application (R Core Team 2021).

II.4. Statistical analyses

We used logistic regressions to analyse the effect of the selected environmental variables on the probability (p) of observing a threatened species. We analysed forest, edge and non-forest species separately for each taxonomic group. In each case, the parks where the number of observations of threatened species was below five for a taxonomic group were excluded from the analyses of the taxonomic group concerned.

From the 69 126 potential pseudo-absences generated, 10 000 were drawn at random and weighted so that the total weight of pseudo-absences equalled the total weight of presences in each model, as recommended by [START_REF] Barbet-Massin | Selecting pseudo-absences for species distribution models: how, where and how many?[END_REF]. We then fitted and compared three logistic regression models. First, we started with a full model (Mfull) with the following environmental variables as predictors: national park identity, elevation, stand tree species composition, distance to the closest path and slope (see equation 1). We calculated the difference in AIC (ΔAIC) between each model and the model MEnv and estimated the values of the coefficients (βi and β'i) associated with historical and current forest area in each model. We used the coefficient of the model with the lowest AIC for interpretation.

To ensure that pseudo-absence random-draws had no effect, we repeated this procedure 20 times using 20 random draws of 10 000 pseudo-absences and results were summarised by their mean ± standard deviation. The coefficients for each buffer size are provided in Appendix S2.

III. Results

III.1. Data summary

Of the 8 061 species recorded, 696 were identified as threatened. Spermatophyta, Coleoptera and Lepidoptera were the most diverse groups while Spermatophyta and Aves had the largest number of observations. All three habitat preferences (forest species: FS, edge species: ES and non-forest species: NFS) were present in the different taxonomic groups, except for Squamata, Amphibia, Odonata and Orthoptera, which contained only non-forest species (see the list of threatened species in Appendix S3).

III.2. Response of threatened species to current or historical forest area in the landscape

In accordance with our first hypothesis, the probability of observing a threatened forest species rose with increasing current forested area in the landscape while non-forest species were negatively affected (Figure 16 and Appendix S2). Current forested area had a significant positive effect on threatened FS spermatophytes (β750 = 2.6 ± 0.1), birds (β1000= 1.2 ± 0.08) and beetles (β250= 5.4 ± 0.06). Conversely, NFS spermatophytes (β100 = -2.8 ± 0.1) and birds (β250 = -2.7 ± 0.1) as well as ES lepidopterans (β1500 = -3.5 ± 0.07) were negatively affected by the current forested area in the landscape.

Our second hypothesis was also verified for several taxonomic groups. Some groups of threatened forest species responded to landscape changes with delay and their presence was better explained by the historical landscape than by the current one (Figure 1). FS Spermatophytes (β1500 = 1.5 ± 0.08), bryophytes (β500 = 0.8 ± 0.04) and beetles (β250 = 2.0 ± 0.02), as well as ES pteridophytes (β250 = 1.4 ± 0.03) were positively affected by the amount of historical forest in the landscape, Mhist being a better model than MAct. In addition, historical forested area negatively affected ES lepidopterans (β100 = -1.7 ± 0.03).

Neither historical nor current forested area in the landscape affected odonatans, Orthopterans, chiropterans, reptiles, amphibians, lichens, NFS pteridophytes, NFS and ES bryophytes (Figure 1).

Results were robust to pseudo-absence random-draws, ΔAIC varied very little and all the β coefficients were included into the IC95% calculated by the first model of the 20 iterations of pseudoabsence random-draws (Appendix S2).

The optimal landscape scale of effect varied among taxonomic groups and the effect tested (current or historical forested area). Historical forest affected bryophytes, beetles and lepidopterans at a rather small landscape scale, whereas spermatophytes were influenced at a large landscape scale. On the other hand, current forest area affected spermatophytes and coleopterans at a small landscape scale while birds and lepidopterans were influenced at a large landscape scale (Figure 1).

IV. Discussion

IV.1. Effect of historical and current landscape context on the probability of observing a threatened species

Several forest taxonomic groups responded positively to the amount of current forest in the landscape. These results confirm our first hypothesis and show the importance of preserving habitats for the maintenance of threatened species [START_REF] Pykälä | Habitat loss and deterioration explain the disappearance of populations of threatened vascular plants, bryophytes and lichens in a hemiboreal landscape[END_REF]. However, FS chiropterans did not respond to current forest cover but this lack of effects for chiropterans may be due to the small number of observations that may have led to a limited statistical power of the models (N=24). FS bryophytes did not respond to current forest cover but this group include both epiphytic and non-epiphytic species that may be poorly affected by the amount of forest in the landscapes [START_REF] Mccune | Comparing the effect of landscape context on vascular plant and bryophyte communities in a human-dominated landscape[END_REF][START_REF] Nordén | Effects of ecological continuity on species richness and composition in forests and woodlands: A review[END_REF]. Conversely, current forested area negatively affected NFS spermatophytes and birds with a small landscape scale of effect, suggesting these species can be found in forests, but they might not tolerate dense forest cover. More investigation is needed, mixing observations inside and outside forest habitats, to properly analyse the effect of land abandonment and afforestation on threatened non-forest species.

Historical forested area better explained the presence of threatened forest spermatophytes than did current forested area at the largest landscapes scale, indicating that these species can be better maintained in large historically wooded areas, i.e. probably the areas least influenced by human activities. Indeed, forest cores may have been less affected by wood and litter extraction or by grazing, which led to the degradation of the forest cover during the 20 th century [START_REF] Jalut | Histoire des forêts du versant nord des Pyrénées au cours des 30000 dernières années[END_REF][START_REF] Leroy | L'histoire des forêts des alpes[END_REF][START_REF] Tochon | Chapitre VII: Les forêts de la Savoie au XVIII° siècle, in: Histoire de l'agriculture en Savoie[END_REF]. Our results agree with [START_REF] Lavergne | Environmental and human factors influencing rare plant local occurrence, extinction and persistence: a 115-year study in the Mediterranean region[END_REF], who showed that rare species are maintained in the least anthropogenised ecosystems, and with [START_REF] Kimberley | Traits of plant communities in fragmented forests: the relative influence of habitat spatial configuration and local abiotic conditions[END_REF], who showed that AFs support more rare species than RFs when AF patches are large. [START_REF] Flensted | Red-listed species and forest continuity -A multi-taxon approach to conservation in temperate forests[END_REF] showed that the richness of red-listed spermatophytes was better correlated to former forested area than to the current one on 10x10km grids, as did [START_REF] Paltto | At which spatial and temporal scales does landscape context affect local density of Red Data Book and Indicator species?[END_REF] in Sweden with 5-km-radius circles. These studies, as well as our own results, suggest that i) the optimal landscape scale of effect is certainly greater than the 1.5-km radius covered by our largest buffer and ii) that the effect occurs in forest landscapes with various degrees of forest fragmentation.

ES pteridophytes were also positively affected by historical forest area while current forest had no effect. However, this taxonomic group was dominated by Dryopteris oreades and Dryopteris ardechensis wich are mostly found in screes, an unfavourable habitat for cultivation that is usually found in ancient forests, which may indirectly explain our result.

Current forest area had a greater impact on FS and NFS birds than historical forest area. These species are highly mobile and are probably not limited by their dispersion ability, which explains the large landscape scale effect for current forest area. High forest cover is beneficial for FS birds but has a significant negative effect on NFS birds and management should be adapted according to conservation objectives [START_REF] Ram | Forest clear-cuts as habitat for farmland birds and butterflies[END_REF].

The stronger effect of historical than current landscape on threatened forest beetles is in agreement with Flensted et al.'s (2016) results. Most of the threatened forest beetles we analysed were actually saproxylic beetles. These species are highly dependent on the forest context and may have a limited dispersal ability [START_REF] Irmler | Species richness of saproxylic beetles in woodlands is affected by dispersion ability of species, age and stand size[END_REF], which could induce a colonization credit in RFs [START_REF] Brin | Effects of forest continuity on flying saproxylic beetle assemblages in small woodlots embedded in agricultural landscapes[END_REF]. However, when the landscape is slightly fragmented, the distribution of saproxylic beetles seems to be more affected by the habitat quality (the abundance and diversity of dead wood, in particular) than by dispersal limitation [START_REF] Janssen | Are biodiversity patterns of saproxylic beetles shaped by habitat limitation or dispersal limitation? A case study in unfragmented montane forests[END_REF]. Furthermore, forest continuity and stand maturity may have additive effects on the species richness and functional composition of saproxylic beetle communities [START_REF] Janssen | Forest continuity acts congruently with stand maturity in structuring the functional composition of saproxylic beetles[END_REF]. If forest continuity is a necessary (but insufficient) condition to obtain very mature stands [START_REF] Nordén | Effects of ecological continuity on species richness and composition in forests and woodlands: A review[END_REF], it is also true that past human societies cleared forests for agriculture and pasture in the most favourable and accessible areas. AFs are thus more frequently found in the least accessible or steepest areas [START_REF] Abadie | Forest recovery since 1860 in a Mediterranean region: drivers and implications for land use and land cover spatial distribution[END_REF][START_REF] Flinn | Environmental causes and consequences of forest clearance and agricultural abandonment in central New York, USA[END_REF][START_REF] Thomas | Long Term Changes in Forest Landscapes in Five National Parks in Metropolitan France and the Future Champagne and Bourgogne National Forest Park[END_REF]. Therefore, AFs may be subject to less logging pressure and thus become more mature than RFs, which could result in an indirect effect of historical forest cover. Our study did not distinguish between these two effects. Further investigation is required to properly disentangle the forest continuity and maturity effects on the response of species, especially in mountain forests [START_REF] Janssen | Do not drop OLD for NEW: conservation needs both forest continuity and stand maturity[END_REF].

Forest bryophytes were also positively affected by the amount of historical forest in the landscape. Because of their low dispersion capacities and their sensitivity to habitat change, bryophytes appear to be good indicator species for forest continuity [START_REF] Mölder | Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)[END_REF] and our results show the importance of ancient forests for the conservation of threatened species. However, the model MHist is only slightly better than MEnv. This weak difference is probably due to the heterogeneity of the FS group that includes both epiphytic and non-epiphytic species able to persist in residual canopy areas in cultural landscapes, which may mitigate our results [START_REF] Fenton | Bryophyte (moss and liverwort) conservation under remnant canopy in managed forests[END_REF].

Historical forested area negatively affected ES lepidopterans at the smallest landscape scale, whereas current forested area had no effect at this landscape scale. This suggests that these species occur in forest but only in recent forest due to an extinction debt. Indeed, past land use can indirectly affect butterfly communities because some grassland species may persist in forests for more than 100 years after canopy closure [START_REF] Burst | Interactive effects of land-use change and distanceto-edge on the distribution of species in plant communities at the forest-grassland interface[END_REF]). Thus, clearcuts in post-agricultural forests have been shown to contain 35% more grassland species than clearcuts in AFs [START_REF] Milberg | Land-use history influence the vegetation in coniferous production forests in southern Sweden[END_REF]. Therefore, vegetation that develops in forest clearcuts on former grasslands favours typical grassland-dependant lepidopterans [START_REF] Ibbe | History matters: Impact of historical land use on butterfly diversity in clear-cuts in a boreal landscape[END_REF], even in clearcuts more than 10 years old [START_REF] Ram | Forest clear-cuts as habitat for farmland birds and butterflies[END_REF]). Our study did not take forest management into account; some lepidopteran observations could have been located in former clearcuts even though the forest was considered closed on the BD FORET ® V2 map (the canopy cover threshold of 40% can be reached in less than 10 years in mountain areas; [START_REF] Fuhr | Large gaps dynamics in the mountain forests of the french Alps[END_REF]. Agricultural land abandonment and tree canopy closure are the two main threats for Lepidoptera [START_REF] Erhardt | Ecology and conservation of alpine Lepidoptera[END_REF][START_REF] Öckinger | Effects of grassland abandonment, restoration and management on butterflies and vascular plants[END_REF]) and our results show that forest-edge Lepidoptera species were negatively affected by the amount of current forest in the landscape at the largest landscape scale. Targeted management in recent forests, for example, maintaining a network of clearings, could be an efficient tool for the conservation of lepidopterans threatened by intensive agricultural practices [START_REF] Ram | Forest clear-cuts as habitat for farmland birds and butterflies[END_REF].

The amount of current forest or the amount of historical forest in the landscape did not affect the other taxonomic groups included in our study. Orthopterans are ectothermic organisms that depend mainly on the amount of heat reaching the ground and are generally negatively affected by forest cover. Thus, the abundance of threatened orthopterans is negatively affected by tree cover and advanced successional stages [START_REF] Helbing | Response of Orthoptera communities to succession in alluvial pine woodlands[END_REF]) and the presence of forest near dry grasslands has a negative effect on their species richness [START_REF] Bieringer | Shading out species richness: edge effect of a pine plantation on the Orthoptera (Tettigoniidae and Acrididae) assemblage of an adjacent dry grassland[END_REF]. Our models probably did not detect any effect due to the low number of observations (n=18) in our dataset. While reptiles are also ectotherms, our results do not show any effect of forested area in the landscape. However, the link between reptile species richness and forested area is unclear in the literature. Indeed, the metaanalysis carried by [START_REF] Thompson | Effects of Secondary Forest Succession on Amphibians and Reptiles: A Review and Meta-analysis[END_REF] does not conclude on differences in species richness or abundance between secondary forests and open areas or between old-growth and secondary forests. In addition, threatened lichens did not respond to either historical or current landscape in our study contrary to Fritz et al.'s (2008) results. However, these authors used a larger continuity gradient (more than 350 years) and used correlation between ancient-forests species richness and threatened species richness. Moreover, lichens are particularly sensitive to the quality of their habitat (substrate, shade, humidity, air pollution, etc.) and maybe not strongly limited by their dispersal ability in slightly fragmented ecosystems [START_REF] Nordén | Effects of ecological continuity on species richness and composition in forests and woodlands: A review[END_REF][START_REF] Janssen | Do not drop OLD for NEW: conservation needs both forest continuity and stand maturity[END_REF]. Finally, odonates may not respond to forest area in our study because they spend most of their life cycle in the larval stage and depend almost exclusively on the quality of the aquatic environment in which they develop. In addition, their high dispersal capacity at the adult stage probably make them rather independent of the landscape context [START_REF] Mcpeek | Chapter 5 : Ecological factors limiting the distributions and abundances of Odonata[END_REF].

IV.2. Use of biodiversity data: limits and perspectives

Like any natural history collection dataset [START_REF] Newbold | Applications and limitations of museum data for conservation and ecology, with particular attention to species distribution models[END_REF], our dataset has several limitations. The first concerns the over-sampling of some taxonomic groups, and even of some species due to societal preferences [START_REF] Troudet | Taxonomic bias in biodiversity data and societal preferences[END_REF]). In our dataset, birds and spermatophytes were better represented than other taxonomic groups, and some charismatic species were more sampled than other species in the same taxonomic group. These species therefore weighted more in our models. The second limitation concerns the spatial distribution of the recorded observations. In the absence of a preplanned sampling design, observers may have selected sites based on previous observations, restricting themselves to easily accessible areas and resampling the same sector over time. Finally, since the sampling protocols, if any, are unknown, the dataset contains no information on species absence. However, despite these weaknesses, these types of databases also has several advantages: they cover large areas and provide information on rare species; they allow a multi-taxonomic approach, which should provide more general results [START_REF] Westgate | Global meta-analysis reveals low consistency of biodiversity congruence relationships[END_REF]) and finally, they can help refine niche models when they are combined with standardised presence/absence data [START_REF] Coron | Bayesian estimation of species relative abundances and habitat preferences using opportunistic data[END_REF] or processed with deep learning techniques [START_REF] Botella | A Deep Learning Approach to Species Distribution Modelling[END_REF]. These types of databases are therefore particularly important tools for ecological research and will become increasingly important with the development of participative science and the era of big data in biodiversity research [START_REF] Hampton | Big data and the future of ecology[END_REF]. We strongly encourage managers to complement biodiversity inventory efforts on taxonomic groups that have been under-sampled because of their low societal interest or due to lack of observers with the necessary scientific skills for species determination.

IV.3. Implications for conservation

In accordance with previous works [START_REF] Flensted | Red-listed species and forest continuity -A multi-taxon approach to conservation in temperate forests[END_REF], our results show that forest continuity matters for red-listed species in landscapes with limited forest fragmentation. They confirm the importance of ancient forests for red-listed species and we therefore encourage policy makers to follow the IUCN recommendations (IUCN 2016) by adopting protective measures for AFs in order to preserve biodiversity.

The current forest area had less effect than the historical forest area (or had no effect) on threatened forest species for some taxonomic groups. On the other hand, high current forest area had a negative effect on some threatened non-forest species. This result questions the effectiveness of afforestation measures to preserve both forest and non-forest red-listed species. Indeed, while afforestation appears to be an effective measure for preserving forest species [START_REF] Newmark | Targeted habitat restoration can reduce extinction rates in fragmented forests[END_REF], the objective could be missed due to the colonisation credit of threatened forest species in recent forests while being detrimental to non-forest threatened species. Landscape planning should be adapted to the conservation objectives but protecting ancient forests is a good compromise to preserve both forest and non-forest threatened species and can be more effective and less expensive than untargeted reforestations.

In addition, plantations and intensive management may also induce loss of forest biodiversity [START_REF] Bremer | Does plantation forestry restore biodiversity or create green deserts? A synthesis of the effects of land-use transitions on plant species richness[END_REF][START_REF] Paillet | Biodiversity Differences between Managed and Unmanaged Forests: Meta-Analysis of Species Richness in Europe[END_REF], which could mitigate differences between recent and ancient forests. Indeed, plantation may cause a recruitement limitation of threatened species in recent forests while intensive management of ancient forest can lead to the loss of their conservation value attributes [START_REF] Bergès | Response of understorey plant communities and traits to past land use and coniferous plantation[END_REF][START_REF] Depauw | Interactive effects of past land use and recent forest management on the understorey community in temperate oak forests in South Sweden[END_REF]. The protection of ancient forests is thus not only about fighting deforestation, but also about sustainable forest management and further research is needed in this way [START_REF] Bergès | Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case[END_REF]. Some authors argue that forest continuity matters more than tree age for conservation of biodiversity [START_REF] Mcmullin | Out with OLD growth, in with ecological continNEWity: new perspectives on forest conservation[END_REF], however, we agree with Janssen et al. 2019 that forest continuity and stand maturity are complementary components because they affect biodiversity by different ways: ancientness is related to species dispersal limitations while stand maturity is related to habitat requirements [START_REF] Janssen | Forest continuity acts congruently with stand maturity in structuring the functional composition of saproxylic beetles[END_REF]). Nevertheless, ancient forests are easier to map than mature forests and managers should use historical maps to identify new high-priority conservation areas.

V. Conclusion

Our results confirm the role of temporal forest continuity in maintaining threatened forest species for multiple taxonomic groups in mountain ecosystems and highlight that the landscape scale of effect differ dramaticaly among taxonomic groups. Our results also show that for some group current forest cover rate is detrimental to non-forest threatened species. Thus, conservation measures will need to strike a balance between the conservation of both groups of species. We encourage stakeholders to prioritise ancient forest conservation measures over afforestation measures to preserve threatened forest species with the least impact on non-forest species.

Figure 1 :

 1 Figure 1: Differences in AIC between models MAct or MHist and MEnv for each taxonomic group and type of threatened species (forest, edge or non-forest). ΔAIC between -2 and +2 (shaded area) means there is no statistical difference between models MAct or MHist and MEnv. Error bars represent standard deviation between the 20 iterations. The higher the ΔAIC, the more explanatory the model. Empty boxes = non-applicable model.

Table 1 :

 1 Description of the forest context in the five National Parks of our study area. Surface areas are in km² and percentages in brackets for current and historical forest area are relative to the total surface area while the historical forest loss is relative to the historical forest area. Forest expansion is relative to the historical forest area, and the share of ancient, deciduous, coniferous and mixed forests are relative to the current forest area.

	National parks		Vanoise	Ecrins	Mercantour	Cevennes	Pyrenees
	Total surface area		1 577	1 702	2 134	3 731	2 523
	Current forest area	298.1 (19%)	571.8 (34%)	1 122.1 (52%)	2 523.2 (68%)	740.6 (29%)
	Historical forest area (1853-1864)	151.3 (10%)	179.8 (11%)	374.9 (18%)	564 (15%)	392.6 (16%)
	Historical forest loss since mid-19 th century	23.7 (16%)	29 (16%)	27.4 (7%)	31.5 (5.6%)	52.8 (13%)
	Forest expansion since mid-19 th century	+ 97%	+ 218%	+ 199%	+ 347%	+ 89%
		Share of ancient forest	43%	26%	31%	21%	46%
		Share of decidous forest	16%	25%	16%	44%	59%
	Current forest	Share of coniferous forest	76%	54%	71%	35%	22%
		Share of mixed forest	8%	21%	13%	21%	19%
		Elevation range (m)	[636 ; 2 480]	[670 ; 2 443]	[153 ; 2 728]	[104 ; 1 698]	[293 ; 2 555]

Appendix S3 :

 S3 List of threatened species classified by taxonomic group with species' forest affinity (FS: forest species; ES: edge species; NFS: non-forest species) and number of occurrences. Ranunculus trichophyllus subsp. eradicatus (Laest.) C.D.K.Cook, 1967 ES Neoorthocaulis floerkei (F.Weber & D.Mohr) L.Söderstr., De Roo & Hedd., 2010 NFS

	Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species Species	Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity Forest affinity	Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences Number occurrences	of of of of of of of of of of of of of of of of of of of of of of of
	Species Bupleurum longifolium L., 1753 Daphne cneorum L., 1753 Hyssopus officinalis L., 1753 Myricaria germanica (L.) Desv., 1824 Veronica spicata L., 1753 Brachythecium tenuicaule (Spruce) Kindb., 1900 Didymodon spadiceus (Mitt.) Limpr., 1888 Heterocladiella dimorpha (Brid.) Ignatov & Fedosov, 2019 Pohlia sphagnicola (Bruch & Schimp.) Broth., 1903 Scapania compacta (Roth) Dumort., 1835 Lichens Chaenothecopsis savonica (Räsänen) Tibell, 1984 Massalongia carnosa (Dicks.) Körb., 1855 Sticta limbata (Sm.) Ach., 1803 Alectoris rufa (Linnaeus, 1758) Glaucidium passerinum (Linnaeus, 1758) Sylvia borin (Boddaert, 1783) Coleoptera Philothermus evanescens (Reitter, 1876) Lysandra hispana (Herrich-Schäffer, 1852) Gampsocleis glabra (Herbst, 1786)	affinity Forest ES ES NFS ES NFS FS ES NFS NFS ES FS ES FS NFS FS NFS FS NFS NFS	occurrences Number 14	of
	Bupleurum rotundifolium L., 1753 Daphne striata Tratt., 1814 Iberis carnosa Willd., 1800 Rhaponticum centauroides (L.) O.Bolòs, 1970 Vicia cassubica L., 1753 Didymodon tophaceus (Brid.) Lisa, 1837 Scapania cuspiduligera (Nees) Müll.Frib., 1915 Acarospora admissa (Nyl.) Kullh., 1871 Chaenothecopsis subparoica (Nyl.) Tibell, 1995 Megalospora tuberculosa (Fée) Sipman, 1983 Sticta sylvatica (Huds.) Ach., 1803 Anser anser (Linnaeus, 1758) Grus grus (Linnaeus, 1758) Sylvia curruca (Linnaeus, 1758) Aeletes atomarius (Aubé, 1843) Phloeostichus denticollis W. Redtenbacher, 1842 Melitaea diamina (Lang, 1789) Polysarcus denticauda (Charpentier, 1825)	NFS NFS ES NFS ES NFS NFS NFS ES FS FS NFS NFS NFS FS FS ES NFS	1
	Calamagrostis pseudophragmites (Haller f.) Koeler, 1802 Dasiphora fruticosa (L.) Rydb., 1898 Illecebrum verticillatum L., 1753 Rhynchospora alba (L.) Vahl, 1805 Vicia disperma DC., 1813 Douinia ovata (Dicks.) H.Buch, 1928 Scapania paludicola Loeske & Müll.Frib., 1915 Acrocordia salweyi (Leight. ex Nyl.) A. L. Sm., 1911 Cladonia sulphurina (Michx.) Fr., 1831 Melanelia panniformis (Nyl.) Essl., 1978 Strigula angustata Cl.Roux & Sérus. Anthus campestris (Linnaeus, 1758) Hirundo rustica Linnaeus, 1758 Sylvia undata (Boddaert, 1783) Ampedus brunnicornis Germar, 1844 Platysoma lineare Erichson, 1834 Nymphalis antiopa (Linnaeus, 1758) Polysarcus scutatus (Brunner von Wattenwyl, 1882)	ES NFS NFS NFS NFS FS NFS FS FS ES FS NFS NFS NFS FS FS ES NFS	1
	Carduus personata (L.) Jacq., 1776 Delphinium consolida L., 1753 Impatiens noli-tangere L., 1753 Ruscus hypoglossum L., 1753 Viola pinnata L., 1753 Encalypta vulgaris Hedw., 1801 Scapania scandica (Arnell & H.Buch) Macvicar, 1926 Ainoa mooreana (Carroll) Lumbsch & I. Schmitt, 2001 Cliostomum corrugatum (Ach.) Fr., 1845 Melanelia tominii (Oksner) Essl. Strigula buxi Chodat, 1912 Anthus pratensis (Linnaeus, 1758) Jynx torquilla Linnaeus, 1758 Tachymarptis melba (Linnaeus, 1758) Anthaxia istriana Rosenhauer, 1847 Prostomis mandibularis (Fabricius, 1801) Papilio alexanor Esper, 1800 Pseudochorthippus montanus (Charpentier, 1825)	NFS NFS ES FS ES ES NFS NFS ES NFS FS NFS NFS NFS FS FS NFS NFS	2
	Carex depauperata Curtis ex With., 1787 Dichoropetalum carvifolia (Vill.) Pimenov & Kljuykov, 2007 Inula bifrons (L.) L., 1763 Salix daphnoides Vill., 1779 Viscaria vulgaris Bernh., 1800 Entosthodon attenuatus (Dicks.) Bryhn, 1908 Scapania umbrosa (Schrad.) Dumort., 1835 Alyxoria ochrocheila (Nyl.) Ertz & Tehler, 2011 Collema glebulentum (Nyl. ex Cromb.) Degel., 1952 Menegazzia terebrata (Hoffm.) A.Massal., 1854 Strigula endolithea Cl.Roux & Bricaud Ardea alba Linnaeus, 1758 Lagopus muta (Montin, 1776) Tetrao urogallus aquitanicus Ingram, 1915 Brachygonus ruficeps (Mulsant & Guillebeau, 1855) Pycnomerus terebrans (Olivier, 1790) Parnassius apollo (Linnaeus, 1758)	FS NFS ES ES NFS ES FS FS NFS ES ES NFS NFS FS FS FS NFS
	Carex depressa Link, 1800 Draba incana L., 1753 Iris graminea L., 1753 Salvia aethiopis L., 1753 Pteridophyta Entosthodon fascicularis (Hedw.) Müll.Hal., 1848 Orthotrichum rogeri Brid., 1812 Schistochilopsis opacifolia (Culm. ex Meyl.) Konstant., 1994 Anaptychia crinalis (Schleich.) Vezda Cyphelium inquinans (Sm.) Trevis., 1862 Micarea adnata Coppins, 1983 Strigula minor (Vezda) Cl.Roux & Sérus. Ardea purpurea Linnaeus, 1766 Lanius senator Linnaeus, 1758 Tetrao urogallus Linnaeus, 1758 Buprestis haemorrhoidalis Herbst, 1780 Pytho depressus Linnaeus, 1767 Parnassius mnemosyne (Linnaeus, 1758)	FS NFS ES ES NFS ES NFS FS FS FS FS NFS NFS FS FS FS ES
	Carex flava L., 1753 Draba nemorosa L., 1753 Juncus capitatus Weigel, 1772 Saussurea alpina (L.) DC., 1810 Allosorus hispanicus (Mett.) Christenh., 2012 Entosthodon muhlenbergii (Turner) Fife, 1985 Orthotrichum scanicum Grönvall, 1885 Schistostega pennata (Hedw.) F.Weber & D.Mohr, 1803 Anema moedlingense Zahlbr., 1908 Cyphelium karelicum (Vain.) Räsänen, 1939 Micarea lutulata (Nyl.) Coppins, 1980 Thelidium dionantense (Hue) Zschacke Ardeola ralloides (Scopoli, 1769) Linaria cannabina (Linnaeus, 1758) Tichodroma muraria (Linnaeus, 1766) Calopus serraticornis (Linnaeus, 1758) Rhizophagus brancsiki Reitter, 1905 Phengaris alcon (Denis & Schiffermüller, 1775)	NFS NFS NFS NFS NFS NFS ES FS NFS FS NFS NFS NFS NFS NFS FS FS NFS
	Carex hordeistichos Vill., 1779 Dracocephalum austriacum L., 1753 Juniperus phoenicea L., 1753 Saxifraga cuneifolia subsp. cuneifolia L., 1759 Allosorus tinaei (Tod.) Christenh., 2012 Entosthodon obtusus (Hedw.) Lindb., 1865 Orthotrichum stellatum Brid., 1826 Sciuro-hypnum curtum (Lindb.) Ignatov, 2008 Arthonia leucopellaea (Ach.) Almq., 1880 Cyphelium notarisii (Tul.) Blomb. & Forssell, 1880 Microcalicium disseminatum (Ach.) Vain., 1927 Thelotrema lepadinum (Ach.) Ach., 1803 Bonasa bonasia (Linnaeus, 1758) Merops apiaster Linnaeus, 1758 Turdus pilaris Linnaeus, 1758 Cardiophorus anticus Erichson, 1840 Rosalia alpina (Linnaeus, 1758) Pieris ergane (Geyer, 1828)	NFS NFS ES FS NFS NFS ES FS FS ES FS FS FS NFS NFS FS FS NFS
	Carex limosa L., 1753 Drosera rotundifolia L., 1753 Kalmia procumbens (L.) Gift, Kron & P.F.Stevens ex Galasso, Banfi & F.Conti, 2005 Saxifraga hirculus L., 1753 Botrychium matricariifolium (Retz.) W.D.J.Koch, 1845 Entosthodon pulchellus (H.Philib.) Brugués Orthotrichum tenellum Bruch ex Brid., 1827 Ptychostomum elegans (Nees) D.Bell & Holyoak, 2020 Scleropodium touretii (Brid.) L.F.Koch, 1949 Aspicilia laevata (Ach.) Arnold, 1887 Cyphelium pinicola Tibell, 1969 Neocatapyrenium radicescens (Nyl.) Breuss, 1996 Trapelia placodioides Coppins & P. James, 1984 Bubo bubo (Linnaeus, 1758) Monticola saxatilis (Linnaeus, 1766) Turdus torquatus Linnaeus, 1758 Cerambyx cerdo Linnaeus, 1758 Sphaerites glabratus (Fabricius, 1792) Polygonia egea (Cramer, 1775)	NFS NFS NFS NFS NFS NFS NFS ES ES ES ES NFS ES NFS NFS FS FS FS NFS
	Carex mairei Coss. & Germ., 1840 Epipogium aphyllum Sw., 1814 Lamium galeobdolon subsp. montanum (Pers.) Hayek, 1929 Saxifraga iratiana F.W.Schultz, 1851 Cystopteris dickieana R.Sim, 1848 Eucladium verticillatum (With.) Bruch & Schimp., 1846 Oxyrrhynchium schleicheri (R.Hedw.) Röll Ptychostomum imbricatulum (Müll.Hal.) Holyoak & N.Pedersen, 2007 Scorpidium cossonii (Schimp.) Hedenäs, 1989 Bacidina caligans (Nyl.) P. Clerc Degelia atlantica (Degel.) P. M. Jørg. & P. James Ochrolechia szatalaensis Vers. Usnea longissima Ach., 1810 Bubulcus ibis (Linnaeus, 1758) Muscicapa striata (Pallas, 1764) Tyto alba (Scopoli, 1769) Cerambyx welensii (Küster, 1845) Stephanopachys quadricollis (Marseul, 1878) Polyommatus amandus (Schneider, 1792)	NFS FS FS NFS NFS ES FS NFS NFS ES FS FS FS NFS NFS NFS FS FS ES
	Carex maritima Gunnerus, 1772 Eriophorum gracile Koch ex Roth, 1806 Lappula squarrosa (Retz.) Dumort., 1827 Schoenus ferrugineus L., 1753 Diphasiastrum alpinum (L.) Holub, 1975 Eurhynchium angustirete (Broth.) T.J.Kop., 1967 Palustriella commutata (Hedw.) Ochyra, 1989 Ptychostomum torquescens (Bruch & Schimp.) Ros & Mazimpaka, 2013 Scorpiurium circinatum (Brid.) M.Fleisch. & Loeske, 1907 Baeomyces placophyllus Ach., 1803 Fuscidea austera (Nyl.) P. James, 1980 Opegrapha vulpina Vondrák, Kocourk. & Tretiach, 2008 Usnea schadenbergiana Göp. & Stein Carduelis carduelis (Linnaeus, 1758) Nucifraga caryocatactes (Linnaeus, 1758) Upupa epops Linnaeus, 1758 Cetonischema speciosissima (Scopoli, 1786) Teredus cylindricus (Olivier, 1790) Polyommatus damon (Denis & Schiffermüller, 1775)	NFS NFS ES NFS ES FS ES ES ES NFS NFS NFS ES NFS FS NFS FS FS NFS
	Carex oedipostyla Duval-Jouve, 1870 Eryngium alpinum L., 1753 Laserpitium gallicum L., 1753 Scrophularia pyrenaica Benth., 1846 Dryopteris ardechensis Fraser-Jenk., 1981 Fissidens exilis Hedw., 1801 Palustriella decipiens (De Not.) Ochyra, 1989 Ptychostomum zieri (Hedw.) Holyoak & N.Pedersen, 2007 Seligeria donniana (Sm.) Müll.Hal., 1848 Bellemerea diamarta (Ach.) Hafellner & Cl.Roux Fuscidea gothoburgensis (H. Magn.) V. Wirth & Vezda Parmeliella testacea P. M. Jørg. Verrucaria constricta Zschacke, 1933 Carduelis citrinella (Pallas, 1764) Nycticorax nycticorax (Linnaeus, 1758) Vanellus vanellus (Linnaeus, 1758) Corticeus longulus (Gyllenhal, 1827) Triplax lacordairii Crotch, 1870 Polyommatus daphnis (Denis & Schiffermüller, 1775)	NFS NFS ES NFS ES ES NFS NFS FS NFS ES FS NFS FS NFS NFS FS FS ES
	Carex olbiensis Jord., 1846 Euphorbia illirica Lam., 1788 Lathraea squamaria L., 1753 Silene inaperta L., 1753 Dryopteris oreades Fomin, 1911 Fissidens gracilifolius Brugg.-Nann. & Nyholm, 1986 Pedinophyllum interruptum (Nees) Kaal., 1893 Racomitrium ericoides (Brid.) Brid., 1819 Seligeria pusilla (Hedw.) Bruch & Schimp., 1846 Bellemerea sanguinea (Kremp.) Hafellner & Cl.Roux Fuscopannaria sampaiana Tav. Peltigera neopolydactyla (Gyeln.) Gyeln., 1932 Verrucaria geophila Zahlbr. Ciconia ciconia (Linnaeus, 1758) Otus scops (Linnaeus, 1758) Squamata Dolotarsus lividus (C.R. Sahlberg, 1833) Triplax melanocephala (Latreille, 1804) Polyommatus dolus (Hübner, 1823)	NFS NFS FS NFS ES FS FS ES FS ES FS ES ES NFS NFS FS FS ES
	Carex pauciflora Lightf., 1777 Euphorbia seguieriana subsp. loiseleurii (Rouy) P.Fourn., 1936 Lathyrus cirrhosus Ser., 1825 Silene noctiflora L., 1753 Lycopodiella inundata (L.) Holub, 1964 Fissidens osmundoides Hedw., 1801 Phaeoceros carolinianus (Michx.) Prosk., 1951 Racomitrium sudeticum (Funck) Bruch & Schimp., 1845 Serpoleskea confervoides (Brid.) Loeske, 1904 Biatora chrysantha (Zahlbr.) Printzen, 1994 Graphis betulina (Pers.) Ach. Peltigera scabrosa Th.Fr., 1861 Verrucula polycarparia Nav.-Ros. & Cl.Roux Ciconia nigra (Linnaeus, 1758) Pandion haliaetus (Linnaeus, 1758) Anguis fragilis Linnaeus, 1758 Elater ferrugineus Linnaeus, 1758 Wanachia triguttata (Gyllenhal, 1810) Polyommatus eros (Ochsenheimer, 1808)	NFS ES ES NFS NFS NFS NFS NFS NFS FS ES NFS ES FS NFS NFS FS FS NFS
	Carlina biebersteinii Bernh. ex Hornem., 1819 Festuca airoides Lam., 1788 Lepidium villarsii Gren. & Godr., 1847 Silene viridiflora L., 1762 Spinulum annotinum (L.) A.Haines, 2003 Fissidens rivularis (Spruce) Schimp., 1851 Phaeoceros laevis (L.) Prosk., 1951 Rhizomnium pseudopunctatum (Bruch & Schimp.) T.J.Kop., 1968 Solenostoma confertissimum (Nees) Schljakov, 1981 Biatora nylanderi Anzi, 1860 Gyalecta derivata (Nyl.) H.Olivier, 1911 Pertusaria glomerata (Ach.) Schaer., 1826 Xylographa trunciseda (Th.Fr.) Minks. ex Redinger Columba livia Gmelin, 1789 Petronia petronia (Linnaeus, 1766) Coronella austriaca Laurenti, 1768 Episernus striatellus (Brisout de Barneville in Grenier, 1863) Xylita laevigata (Hellenius, 1786) Satyrus actaea (Esper, 1781)	NFS NFS NFS ES ES NFS NFS ES NFS FS FS ES FS NFS NFS NFS FS FS NFS
	Circaea alpina L., 1753 Festuca amethystina L., 1753 Linaria angustissima (Loisel.) Borbás, 1900 Spergula segetalis (L.) Vill., 1789 Bryophyta Fossombronia angulosa (Dicks.) Raddi, 1818 Philonotis calcarea (Bruch & Schimp.) Schimp., 1856 Rhynchostegiella curviseta (Brid.) Limpr., 1896 Solenostoma hyalinum (Lyell) Mitt., 1870 Biatora ocelliformis (Nyl.) Arnold, 1870 Hypocenomyce anthracophila (Nyl.) P. James & Gotth. Schneid. Physcia phaea (Tuck.) Thoms. Chiroptera Columba oenas Linnaeus, 1758 Phylloscopus sibilatrix (Bechstein, 1793) Emys orbicularis (Linnaeus, 1758) Erotides cosnardi (Chevrolat, 1831) Zilora obscura (Fabricius, 1794) Satyrus ferula (Fabricius, 1793)	FS NFS NFS NFS NFS NFS ES ES FS ES NFS FS FS NFS FS FS NFS
	Cirsium carniolicum subsp. rufescens (Ramond ex DC.) P.Fourn., 1940 Festuca longifolia Thuill., 1799 Linaria pelisseriana (L.) Mill., 1768 Spiranthes aestivalis (Poir.) Rich., 1817 Amblyodon dealbatus (Hedw.) P.Beauv., 1804 Fossombronia caespitiformis (Raddi) De Not. ex Rabenh., 1860 Philonotis capillaris Lindb., 1867 Rhynchostegium murale (Hedw.) Schimp., 1852 Solenostoma sphaerocarpum (Hook.) Steph., 1901 Biatoridium monasteriense J. Lahm ex Körb., 1860 Hypogymnia austerodes (Nyl.) Räsänen, 1943 Placopsis gelida (L.) Linds., 1866 Miniopterus schreibersii (Natterer in Kuhl, 1817) Corvus corax Linnaeus, 1758 Poecile montanus (Conrad von Baldenstein, 1827) Mauremys leprosa (Schweigger, 1812) Eupotosia mirifica (Mulsant, 1842) Lepidoptera Scolitantides orion (Pallas, 1771)	NFS NFS NFS NFS NFS ES NFS ES NFS FS ES NFS NFS NFS FS NFS FS NFS
	Cirsium glabrum DC., 1815 Fritillaria moggridgei Baker, 1879 Linaria supina (L.) Chaz., 1790 Stipa pennata L., 1753 Andreaea rothii subsp. falcata (Schimp.) Lindb., 1879 Fossombronia pusilla (L.) Nees, 1838 Philonotis rigida Brid., 1827 Rhytidiadelphus subpinnatus (Lindb.) T.J.Kop., 1971 Southbya tophacea (Spruce) Spruce, 1850 Bryonora rhypariza (Nyl.) Poelt, 1983 Ionaspis obtecta (Vain.) R. Sant., 2004 Placynthium lismorense (Cromb.) Vain., 1909 Myotis bechsteinii (Kuhl, 1817) Coturnix coturnix (Linnaeus, 1758) Prunella collaris (Scopoli, 1769) Natrix maura (Linnaeus, 1758) Gnorimus variabilis (Linnaeus, 1758) Aricia morronensis (Ribbe, 1910) Zygaena hilaris Ochsenheimer, 1808	NFS NFS NFS NFS NFS NFS NFS FS NFS ES NFS NFS FS NFS NFS NFS FS NFS ES
	Cirsium heterophyllum (L.) Hill, 1768 Gagea bohemica (Zauschn.) Schult. & Schult.f., 1829 Linnaea borealis L., 1753 Streptopus amplexifolius (L.) DC., 1805 Andreaea rothii subsp. rothii F.Weber & D.Mohr, 1807 Fossombronia wondraczekii (Corda) Dumort. ex Lindb., 1873 Philonotis tomentella Molendo, 1864 Riccardia chamedryfolia (With.) Grolle, 1969 Syntrichia montana var. montana Nees, 1819 Bryoria bicolor (Ehrh.) Brodo & D.Hawksw., 1977 Ionaspis suaveolens (Fr.) Th.Fr. ex Stein Porina hoehneliana (Jaap) R. Sant., 1952 Myotis blythii (Tomes, 1857) Delichon urbicum (Linnaeus, 1758) Puffinus mauretanicus P. R. Lowe, 1921 Timon lepidus (Daudin, 1802) Hyperisus declive (Dufour, 1843) Boloria euphrosyne (Linnaeus, 1758) Zygaena trifolii (Esper, 1783)	NFS NFS FS NFS NFS ES NFS ES NFS ES NFS FS NFS NFS NFS NFS FS ES ES
	Cistus laurifolius L., 1753 Galium pusillum L., 1753 Linum austriacum L., 1753 Swertia perennis L., 1753 Anthoceros agrestis Paton, 1979 Fuscocephaloziopsis lunulifolia (Dumort.) Vána & L.Söderstr., 2013 Riccia beyrichiana Hampe, 1838 Syntrichia virescens (De Not.) Ochyra, 1992 Bryoria implexa (Hoffm.) Brodo & D.Hawksw., 1977 Lecanora caesiosora Poelt, 1966 Porina leptosperma Müll.Arg., 1883 Myotis capaccinii (Bonaparte, 1837) Dendrocopos leucotos (Bechstein, 1803) Puffinus puffinus (Brünnich, 1764) Vipera aspis (Linnaeus, 1758) Ischnomera cinerascens (Pandellé in Grénier, 1867) Brenthis ino (Rottemburg, 1775) Zygaena viciae (Denis & Schiffermüller, 1775)	ES NFS NFS NFS ES FS NFS NFS FS ES FS NFS FS NFS NFS FS ES ES
	Cistus umbellatus L., 1753 Galium tricornutum Dandy, 1957 Lunaria rediviva L., 1753 Gongylanthus ericetorum (Raddi) Nees, 1836 Riccia ciliata Hoffm., 1795 Targionia hypophylla L., 1753 Bryoria subcana (Nyl. ex Stizenb.) Brodo & D.Hawksw., 1977 Lecanora eurycarpa Poelt, Leuckert & Cl.Roux non Nyl. Porpidia platycarpoides (Bagl.) Hertel, 1987 Nyctalus lasiopterus (Schreber, 1780) Dendrocopos medius (Linnaeus, 1758) Pyrrhocorax pyrrhocorax (Linnaeus, 1758) Vipera aspis aspis (Linnaeus, 1758) Judolia sexmaculata (Linnaeus, 1758) Carterocephalus palaemon (Pallas, 1771)	ES NFS FS NFS NFS NFS ES NFS NFS FS FS NFS NFS FS ES
	Riccia ciliifera Link ex Lindenb. Thuidium recognitum (Hedw.) Lindb., 1874 Buellia epigaea (Pers.) Tuck. Lecanora orbicularis (Schaer.) Vain., 1899 Dendrocopos minor (Linnaeus, 1758)	NFS FS NFS NFS FS
	Riccia crozalsii Levier, 1902 Timmia austriaca Hedw., 1801 Calicium trabinellum (Ach.) Ach., 1810 Lecidea albohyalina (Nyl.) Th.Fr., 1874 Egretta garzetta (Linnaeus, 1766)	NFS ES FS FS NFS
	Riccia gougetiana Durieu & Mont., 1849 Tortella inclinata (R.Hedw.) Limpr., 1888 Caloplaca conversa (Kremp.) Jatta, 1900 Lecidea paupercula Th.Fr., 1874 Emberiza cia Linnaeus, 1766	NFS NFS NFS NFS NFS
	Riccia nigrella DC., 1815 Tortula atrovirens (Sm.) Lindb., 1864 Caloplaca exsecuta (Nyl.) Dalla Torre & Sarnth., 1902 Lepraria lobificans Nyl., 1873 Emberiza citrinella Linnaeus, 1758	NFS NFS NFS NFS NFS
	Riccia subbifurca Warnst. ex Croz., 1903 Tortula inermis (Brid.) Mont., 1832 Catolechia wahlenbergii (Ach.) Körb., 1855 Leptogium corticola (Taylor) Tuck., 1849 Emberiza hortulana Linnaeus, 1758	FS NFS ES FS NFS
	Riccia warnstorfii Limpr. ex Warnst., 1899 Tortula marginata (Bruch & Schimp.) Spruce, 1845 Chaenotheca brunneola (Ach.) Müll.Arg., 1862 Leptogium furfuraceum (Harm.) Sierk, 1964 Emberiza schoeniclus (Linnaeus, 1758)	NFS ES FS ES NFS
	Saccogyna viticulosa (L.) Dumort., 1831 Trichostomum brachydontium Bruch, 1829 Chaenotheca phaeocephala (Turner) Th.Fr., 1861 Llimoniella phaeophysciae Diederich, Ertz & Etayo, 2010 Falco naumanni Fleischer, 1818	FS ES ES FS NFS
	Saelania glaucescens (Hedw.) Broth., 1894 Trichostomum crispulum Bruch, 1829 Chaenothecopsis epithallina Tibell, 1975 Falco peregrinus Tunstall, 1771	NFS ES FS NFS
	Scapania aequiloba (Schwägr.) Dumort., 1835 Ulota hutchinsiae (Sm.) Hammar, 1852 Chaenothecopsis pusiola (Ach.) Vain., 1927 Ficedula hypoleuca (Pallas, 1764) Orthoptera	ES ES FS FS
	Radiola linoides Roth, 1788 Didymodon glaucus Ryan, 1901 Scapania aspera M.Bernet & Bernet, 1888 Gallinago gallinago (Linnaeus, 1758)	NFS NFS ES NFS

Spermatophyta Achillea erba-rotta subsp. erba-rotta All., 1773 ES Achillea nobilis L., 1753 NFS Aconitum napellus subsp. burnatii (Gáyer) J.-M.Tison, 2010 NFS Adonis aestivalis L., 1762 NFS Adonis flammea Jacq., 1776 NFS Adonis vernalis L., 1753 NFS Aethionema saxatile (L.) W.T.Aiton, 1812 ES Alchemilla plicata Buser, 1893 NFS Anacamptis coriophora (L.) R.M.Bateman, Pridgeon & M.W.Chase, 1997 NFS Anacamptis laxiflora (Lam.) R.M.Bateman, Pridgeon & M.W.Chase, 1997 NFS Androsace septentrionalis L., 1753 NFS Apera interrupta (L.) P.Beauv., 1812 NFS Aphyllanthes monspeliensis L., 1753 NFS Arabis auriculata Lam., 1783 NFS Arenaria ligericina Lecoq & Lamotte, 1847 NFS Arenaria montana L., 1755 ES Artemisia chamaemelifolia Vill., 1779 NFS Asperula tinctoria L., 1753 ES Aster amellus L., 1753 ES Aster pyrenaeus Desf. ex DC., 1805 NFS Astragalus alopecuroides L., 1753 NFS Astragalus penduliflorus Lam., 1779 NFS Astragalus vesicarius subsp. pastellianus (Pollini) Arcang., 1882 NFS Astragalus vesicarius subsp. vesicarius L., 1753 NFS Blitum virgatum L., 1753 ES Brachypodium retusum (Pers.) P.Beauv., 1812 NFS Briza minor L., 1753 NFS Buglossoides incrassata subsp. permixta (Jord.) L.Cecchi & Selvi, 2014 ES Cochlearia pyrenaica DC., 1821 NFS Coeloglossum viride (L.) Hartm., 1820 NFS Colchicum alpinum DC., 1805 NFS Corallorhiza trifida Châtel., 1760 FS Cotoneaster nebrodensis (Guss.) K.Koch, 1853 ES Crocus vernus (L.) Hill, 1765 NFS Cruciata glabra (L.) Ehrend., 1958 ES Cynoglossum germanicum Jacq., 1767 ES Cytinus hypocistis (L.) L., 1767 ES Daboecia cantabrica (Huds.) K.Koch, 1872 ES Gentiana utriculosa L., 1753 NFS Gratiola officinalis L., 1753 NFS Gymnadenia odoratissima (L.) Rich., 1817 NFS Hackelia deflexa (Wahlenb.) Opiz, 1838 ES Hedysarum boutignyanum (A.Camus) Alleiz., 1928 NFS Hedysarum brigantiacum Bourn., Chas & Kerguélen, 1992 NFS Herminium monorchis (L.) R.Br., 1813 NFS Hieracium isolanum (Besse & Zahn) Zahn, 1916 NFS Horminum pyrenaicum L., 1753 NFS Hypochaeris uniflora Vill., 1779 NFS Luzula desvauxii Kunth, 1841 NFS Lysimachia minima (L.) U.Manns & Anderb., 2009 NFS Lysimachia tenella L., 1753 NFS Lythrum hyssopifolia L., 1753 NFS Matthiola valesiaca J.Gay ex Boiss., 1867 NFS Mentha arvensis L., 1753 NFS Micranthes clusii (Gouan) B.Bock, 2012 NFS Moehringia lebrunii Merxm., 1965 NFS Molopospermum peloponnesiacum (L.) W.D.J.Koch, 1824 NFS Muscari botryoides (L.) Mill., 1768 NFS Neotinea maculata (Desf.) Stearn, 1974 NFS Neottia cordata (L.) Rich., 1817 FS Nepeta nuda L., 1753 ES Nothobartsia spicata (Ramond) Bolliger & Molau, 1992 NFS Odontites luteus subsp. lanceolatus (Gaudin) P.Fourn., 1937 NFS Omalotheca norvegica (Gunnerus) Sch.Bip. & F.W.Schultz, 1861 ES Ophrys speculum Link, 1799 NFS Orobanche rapum-genistae Thuill., 1799 ES Orobanche salviae F.W.Schultz ex W.D.J.Koch, 1833 FS Oxytropis fetida (Vill.) DC., 1802 NFS Oxytropis pilosa (L.) DC., 1802 NFS Phelipanche arenaria (Borkh.) Pomel, 1874 NFS Phelipanche lavandulacea (F.W.Schultz) Pomel, 1874 NFS Phyteuma cordatum Balb., 1809 NFS Pinguicula alpina L., 1753 NFS Pinguicula grandiflora subsp. rosea (Mutel) Casper, 1962 NFS Pinguicula longifolia subsp. longifolia Ramond ex DC., 1805 NFS Pinguicula reichenbachiana Schindl., 1908 NFS Pinus nigra subsp. salzmannii (Dunal) Franco, 1943 FS Poa hybrida Gaudin, 1808 NFS Polemonium caeruleum L., 1753 NFS Polygala alpina (DC.) Steud., 1821 NFS Potamogeton natans L., 1753 ES Potentilla alba L., 1753 ES Potentilla cinerea Chaix ex Vill., 1779 NFS Potentilla delphinensis Gren. & Godr., 1848 NFS Potentilla fagineicola Lamotte, 1877 NFS Psilurus incurvus (Gouan) Schinz & Thell., 1913 NFS Quercus cerris L., 1753 FS Symphytum bulbosum K.F.Schimp., 1825 ES Tephroseris helenitis (L.) B.Nord., 1978 NFS Thalictrum lucidum L., 1753 ES Trichophorum alpinum (L.) Pers., 1805 NFS Trifolium saxatile All., 1773 NFS Trinia glauca (L.) Dumort., 1827 NFS Trochiscanthes nodiflora (All.) W.D.J.Koch, 1824 ES Turgenia latifolia (L.) Hoffm., 1814 NFS Utricularia minor L., 1753 ES Vaccinium microcarpum (Turcz. ex Rupr.) Schmalh., 1871 ES Veronica montana L., 1755 NFS Atrichum angustatum (Brid.) Bruch & Schimp., 1844 ES Barbilophozia lycopodioides (Wallr.) Loeske, 1907 FS Bartramia stricta Brid., 1803 NFS Bazzania flaccida (Dumort.) Grolle, 1972 FS Biantheridion undulifolium (Nees) Konstant. & Vilnet, 2010 NFS Blasia pusilla L., 1753 NFS Brachydontium trichodes (F.Weber) Milde, 1869 FS Brachytheciastrum velutinum (Hedw.) Ignatov & Huttunen, 2002 FS Brachythecium glareosum (Bruch ex Spruce) Schimp., 1853 ES Brachythecium mildeanum (Schimp.) Schimp., 1862 ES Brachythecium tommasinii (Sendtn. ex Boulay) Ignatov & Huttunen, 2002 NFS Bryum gemmiparum De Not., 1865 NFS Bryum ruderale Crundw. & Nyholm, 1963 NFS Buckia vaucheri (Lesq.) D.Ríos, M.T.Gallego & J.Guerra, 2018 NFS Calypogeia arguta Nees & Mont., 1838 ES Calypogeia suecica (Arnell & J.Perss.) Müll.Frib., 1904 FS Campyliadelphus chrysophyllus (Brid.) R.S.Chopra ES Campylium protensum (Brid.) Kindb., 1894 ES Campylophyllopsis calcarea (Crundw. & Nyholm) Ochyra, 2010 FS Campylopus pilifer Brid., 1819 NFS Cephaloziella baumgartneri Schiffn., 1905 NFS Cephaloziella dentata (Raddi) Steph., 1897 NFS Cephaloziella hampeana (Nees) Schiffn.ex Loeske, 1903 ES Cephaloziella integerrima (Lindb.) Warnst., 1902 FS Cephaloziella phyllacantha (C.Massal. & Carestia) Müll.Frib., 1913 FS Cephaloziella rubella (Nees) Warnst., 1902 FS Cephaloziella stellulifera (Taylor ex Carrington & Pearson) Croz., 1903 NFS Cephaloziella turneri (Hook.) Müll.Frib., 1913 NFS Cinclidotus danubicus Schiffn. & Baumgartner, 1906 NFS Codonoblepharon forsteri (Dicks.) Goffinet, 2004 FS Conardia compacta (Drumm. ex Müll.Hal.) H.Rob. FS Coscinodon cribrosus (Hedw.) Spruce, 1849 NFS Crossocalyx hellerianus (Nees ex Lindenb.) Meyl., 1939 FS Cynodontium strumiferum (Hedw.) Lindb., 1864 ES Dicranum fuscescens Sm. FS Dicranum muehlenbeckii Bruch & Schimp., 1847 FS Dicranum spadiceum J.E.Zetterst., 1865 ES Didymodon acutus (Brid.) K.Saito, 1975 NFS Didymodon ferrugineus (Schimp. ex Besch.) M.O.Hill, 1981 NFS Grimmia elatior Bruch ex Bals.-Criv. & De Not., 1838 NFS Grimmia funalis (Schwägr.) Bruch & Schimp., 1845 NFS Grimmia incurva Schwägr., 1811 NFS Grimmia longirostris Hook., 1818 NFS Grimmia muehlenbeckii Schimp., 1860 ES Grimmia orbicularis Bruch ex Wilson, 1844 NFS Grimmia tergestina Tomm. ex Bruch & Schimp., 1845 NFS Gymnostomum viridulum Brid., 1826 NFS Gyroweisia tenuis (Hedw.) Schimp., 1876 ES Habrodon perpusillus (De Not.) Lindb., 1863 FS Heterocladium wulfsbergii I.Hagen, 1909 FS Homalothecium philippeanum (Spruce) Schimp., 1851 ES Hydrogonium croceum (Brid.) Jan Kucera, 2013 NFS Hygrohypnum luridum (Hedw.) Jenn., 1913 ES Hylocomiastrum umbratum (Hedw.) M.Fleisch. ex Broth., 1925 FS Hyocomium armoricum (Brid.) Wijk & Margad., 1961 ES Isopterygiella pulchella (Hedw.) Ignatov & Ignatova, 2020 NFS Kiaeria starkei (F.Weber & D.Mohr) I.Hagen, 1915 NFS Leptodon smithii (Hedw.) F.Weber & D.Mohr, 1803 FS Lescuraea plicata (Schleich. ex F.Weber & D.Mohr) Broth. NFS Lescuraea radicosa (Mitt.) Mönk., 1927 NFS Lescuraea saxicola (Schimp.) Molendo, 1864 NFS Lewinskya acuminata (H.Philib.) F.Lara, Garilleti & Goffinet, 2016 ES Lewinskya shawii (Wilson) F.Lara, Garilleti & Goffinet, 2016 NFS Lophocolea fragrans (Moris & De Not.) Gottsche, Lindenb. & Nees, 1845 FS Lophozia ascendens (Warnst.) R.M.Schust., 1952 FS Mannia androgyna (L.) A.Evans, 1938 NFS Mannia fragrans (Balb.) Frye & L.Clark, 1937 NFS Mannia triandra (Scop.) Grolle, 1975 NFS Marchantia quadrata Scop., 1772 NFS Meesia uliginosa Hedw., 1801 NFS Mesoptychia bantriensis (Hook.) L.Söderstr. & Vána, 2012 NFS Mesoptychia heterocolpos (Thed. ex Hartm.) L.Söderstr. & Vána, 2012 NFS Mesoptychia turbinata (Raddi) L.Söderstr. & Vána, 2012 NFS Mnium marginatum (Dicks.) P.Beauv. FS Mnium spinosum (Voit) Schwägr., 1816 FS Mnium thomsonii Schimp., 1876 ES Mylia anomala (Hook.) Gray, 1821 NFS Myurella julacea (Schwägr.) Schimp., 1853 NFS Neoorthocaulis attenuatus (Mart.) L.Söderstr., De Roo & Hedd., 2010 ES Obtusifolium obtusum (Lindb.) S.W.Arnell, 1956 FS Odontoschisma fluitans (Nees) L.Söderstr. & Vána, 2013 NFS Orthothecium intricatum (Hartm.) Schimp., 1851 FS Orthotrichum pumilum Sw. ex anon. NFS Plagiomnium elatum (Bruch & Schimp.) T.J.Kop., 1968 NFS Plagiomnium medium (Bruch & Schimp.) T.J.Kop., 1968 FS Plagiomnium rostratum (Schrad.) T.J.Kop., 1968 ES Plagiothecium laetum Schimp., 1851 FS Plagiothecium piliferum (Sw.) Schimp., 1851 NFS Plagiothecium platyphyllum Mönk., 1927 FS Plasteurhynchium striatulum (Spruce) M.Fleisch., 1925 FS Platydictya jungermannioides (Brid.) H.A.Crum, 1964 FS Platyhypnum duriusculum (De Not.) Ochyra, 2013 NFS Pogonatum nanum (Schreb. ex Hedw.) P.Beauv., 1805 ES Pohlia bulbifera (Warnst.) Warnst., 1904 NFS Pohlia proligera (Kindb.) Lindb. ex Broth. ES Polytrichastrum alpinum (Hedw.) G.L.Sm., 1971 ES Porella obtusata (Taylor) Trevis., 1877 NFS Pottiopsis caespitosa (Brid.) Blockeel & A.J.E.Sm. NFS Pterygoneurum ovatum (Hedw.) Dixon, 1934 NFS Ptilidium pulcherrimum (Weber) Vain. FS Ptilium crista-castrensis (Hedw.) De Not., 1867 FS Ptychostomum creberrimum (Taylor) J.R.Spence & H.P.Ramsay, 2005 NFS Lobaria linita (Ach.) Rabenh., 1845 ES Lobaria virens (With.) J. R. Laundon FS Lobothallia parasitica (B. de Lesd.) NFS Pycnora sorophora (Vain.) Hafellner, 2001 ES Ramboldia elabens (Fr.) Kantvilas & Elix, 2007 ES Ramonia subsphaeroides (Tav.) Vezda FS Rhizocarpon postumum (Nyl.) Arnold, 1870 ES Rhizocarpon subgeminatum Eitn. NFS Rosellinula haplospora (Th.Fr. & Almq.) R. Sant., 1986 NFS Sarcogyne fallax H. Magn., 1935 NFS Schaereria cinereorufa (Schaer.) Th.Fr., 1861 ES Solenopsora liparina (Nyl.) Zahlbr., 1919 NFS Sticta fuliginosa (Dicks.) Ach., 1803 FS Nyctalus noctula (Schreber, 1774) FS Rhinolophus euryale Blasius, 1853 ES Rhinolophus ferrumequinum (Schreber, 1774) ES Aves Acrocephalus scirpaceus (Hermann, 1804) NFS Actitis hypoleucos (Linnaeus, 1758) NFS Aegolius funereus (Linnaeus, 1758) FS Alcedo atthis (Linnaeus, 1758) NFS Alectoris graeca (Meisner, 1804) NFS Pyrrhula pyrrhula (Linnaeus, 1758) FS Rallus aquaticus Linnaeus, 1758 NFS Saxicola rubetra (Linnaeus, 1758) NFS Saxicola rubicola (Linnaeus, 1766) NFS Scolopax rusticola Linnaeus, 1758 FS Serinus serinus (Linnaeus, 1766) NFS Spatula clypeata (Linnaeus, 1758) NFS Spinus spinus (Linnaeus, 1758) FS Sternula albifrons (Pallas, 1764) NFS Streptopelia turtur (Linnaeus, 1758) NFS Vipera aspis zinnikeri Kramer, 1958 NFS Vipera berus (Linnaeus, 1758) NFS Amphibia Alytes obstetricans (Laurenti, 1768) NFS Bombina variegata (Linnaeus, 1758) NFS Calotriton asper (Al. Dugès, 1852) NFS Pelophylax kl. esculentus (Linnaeus, 1758) NFS Rana pyrenaica Serra-Cobo, 1993 NFS Triturus marmoratus (Latreille, 1800) NFS Kisanthobia ariasi (Robert, 1858) FS Lamia textor (Linnaeus, 1758) FS Melandrya dubia (Schaller, 1783) FS Merohister ariasi (Marseul, 1864) FS Morimus asper (Sulzer, 1776) FS Mycetochara quadrimaculata (Latreille, 1804) FS Necydalis ulmi Chevrolat, 1838 FS Osmoderma eremita (Scopoli, 1763) FS Oxylaemus variolosus (Dufour, 1843) FS Chazara briseis (Linnaeus, 1764) NFS Coenonympha dorus (Esper, 1782) NFS Cupido minimus (Fuessly, 1775) NFS Erebia gorgone Boisduval, 1833 NFS Erebia pronoe (Esper, 1780) NFS Eumedonia eumedon (Esper, 1780) ES Fabriciana niobe (Linnaeus, 1758) ES Hesperia comma (Linnaeus, 1758) NFS Hyponephele lycaon (Rottemburg, 1775) NFS Odonata Aeshna juncea (Linnaeus, 1758) NFS Coenagrion hastulatum (Charpentier, 1825) NFS Cordulegaster bidentata Selys, 1843 NFS Lestes dryas Kirby, 1890 NFS Lestes sponsa (Hansemann, 1823) NFS Macromia splendens (Pictet, 1843) NFS Sympetrum danae (Sulzer, 1776) NFS Sympetrum flaveolum (Linnaeus, 1758) NFS

OrthopteraSardet, É., Braud, Y., Roesti, C., 2015. Orthoptères de France, Belgique, Luxembourg et Suisse

Acknowledgements

We thank all the National Park managers, National Botanical Conservatories and French Nature and Landscape Information System for sharing their data. We also thank Marie Thomas for coordinating the project and Elodie Le Souchu for her help compining the data. We acknowledge Thomas Legland, Frédéric Archaux and Benoît Renaux for their expertise on different taxonomic groups as well as Vicki Moore for proof reading the manuscript. We also thank the two anonymous rewiewers for their valuable comments.

Appendices

Appendix S1: List of references used to classify species according to their preferred habitat.

Taxonomic groups References

Spermatophyta/ Pteridophyta

Appendix S2: Values of the 20 βi coefficients estimated with different pseudo-absence random draws for the effect of the historical or current forested area according to different buffer sizes.The-shaded area correspond to the 95% confidence interval of the coefficient estimates in the first iteration of pseudo-absence random draws. Empty boxes = non-applicable models. Points for the 20 pseudo-absence random-draws are jittered on the x axis for a better visualisation.