Ronan Marrec
email: ronan.marrec@u-picardie.fr

Vincent Le Roux
email: vincent.leroux@u-picardie.fr

Ludmilla Martin
email: ludmilla.martin0109@gmail.com

Jonathan Roger Michel

Henri Lenoir

Jorg Brunet
email: jorg.brunet@slu.se

Sara A O Cousins
email: sara.cousins@natgeo.su.se

Pallieter De Smedt
email: pallieter.desmedt@ugent.be

Marc Deconchat
email: marc.deconchat@inrae.fr

Martin Diekmann
email: mdiekman@uni-bremen.de

Steffen Ehrmann
email: steffen.ehrmann@idiv.de

Jonathan Lenoir
email: jonathan.lenoir@u-picardie.fr

Jörg Brunet

Emilie Gallet-Moron

Brice Giffard
email: brice.giffard@gmail.com

Jaan Liira
email: jaan.liira@ut.ee

Jessica Lindgren
email: jessica.lindgren@natgeo.su.se

Alicia Valdes
email: aliciavaldes1501@gmail.com

Kris Verheyen
email: kris.verheyen@ugent.be

Monika Wulf
email: mwulf@zalf.de

Guillaume Decocq
email: guillaume.decocq@u-picardie.fr

Jörg Brunet

Multiscale drivers of carabid beetle (Coleoptera: Carabidae) assemblages in small European woodlands

Keywords: Agricultural landscapes, arthropods, biogeographic gradient, carabid beetles, habitat quality, landscape composition, macroclimate, metacommunity dynamics, smallFOREST, species turnover

published or not. The documents may come   L'archive ouverte pluridisciplinaire

 BELSPO (Belgium)as part of the 2011 BiodivERsA call for research proposals. This paper was written while LM held a PhD fellowship from the « Conseil Régional de Picardie ». PDS holds a post-doctoral fellowship of the Research Foundation-Flanders (FWO).

AUTHOR'S CONTRIBUTIONS

GD, KV, MD, VLR, LM, BG, and PDS designed the study. RM, LM, GD, and VLR wrote the manuscript. RM, LM, and JoL ran the analyses. LM, PDS, VLR, and BG sorted arthropods from the traps and identified the species. LM, VLR, AV, JoL, GD, BG, MD, PDS, KV, JB, SC, JaL, MD, AK, MW, TN, and JeL collected data in the field and contributed to write the manuscript.

EGM managed the databases and conducted GIS analyses.

BIOSKETCH

Ronan Marrec is an Assistant Professor at the UMR 7058 CNRS EDYSAN (Ecology and Dynamics of Anthropogenic Systems), Jules Verne University of Picardy, France. As a landscape ecologist he is interested in evaluating the role of the spatial structure and temporal dynamics of landscapes and human activities on biodiversity and ecosystem services. For more information: https://ronanmarrec.weebly.com.

All authors are members of the research project smallFOREST (http://www.upicardie.fr/smallforest/uk/), which is devoted to studying biodiversity and ecosystem services of small forest patches in European landscapes.

INTRODUCTION

The cover of present-day forests in European lowland mainly consists of forest patches of various size, age, tree species composition, and degree of isolation, that are embedded in a more or less intensively managed agricultural matrix [START_REF] Estreguil | Forest landscape in Europe: pattern, fragmentation and connectivity[END_REF]. Forest loss and fragmentation are widely acknowledged as major threats to biodiversity and ecosystem functioning through their effects on habitat availability, quality, and connectivity [START_REF] Fahrig | Effects of habitat fragmentation on biodiversity[END_REF][START_REF] Foley | Global consequences of land use[END_REF][START_REF] Haddad | Habitat fragmentation and its lasting impact on Earth's ecosystems[END_REF]. In particular, increased fragmentation reduces the amount of forest interior habitat which is available for habitat specialist species [START_REF] Pfeifer | Creation of forest edges has a global impact on forest vertebrates[END_REF]. Therefore, patch-level biotic homogenization (i.e., loss of genetic, taxonomic, and functional diversity of species assemblages) due to edge effects is generally observed, which increases in importance as patch size decreases [START_REF] Mckinney | Biotic homogenization: a few winners replacing many losers in the next mass extinction[END_REF][START_REF] Olden | On defining and quantifying biotic homogenization[END_REF]. At the same time, forest edges can offer suitable conditions for a number of animal and plant generalist species originating from neighboring, more open or/and disturbed habitats (e.g., [START_REF] Cousins | The influence of management history and habitat on plant species richness in a rural hemiboreal landscape, Sweden[END_REF][START_REF] González | Natural vegetation cover in the landscape and edge effects: differential responses of insect orders in a fragmented forest[END_REF][START_REF] De Smedt | Linking macrodetritivore distribution to desiccation resistance in small forest fragments embedded in agricultural landscapes in Europe[END_REF], thereby potentially enhancing the delivery of multiple ecosystem services in small forest patches [START_REF] Kütt | Multi-user quality of floral services along a gradient of margin habitats between semi-natural grasslands and forests[END_REF][START_REF] Valdés | High ecosystem service delivery potential of small woodlands in agricultural landscapes[END_REF]. Another consequence of forest loss and fragmentation is that remnant forest patches are increasingly distant from each other. Moreover, the landscape matrix into which forest patches are embedded may be more or less permeable to species movements. Patch isolation and matrix composition both determine landscape connectivity, i.e., the degree to which landscape mosaics facilitate movement of organisms and their genes between distinct forest patches [START_REF] Taylor | Connectivity is a vital element of landscape structure[END_REF][START_REF] Ricotta | Quantifying the network connectivity of landscape mosaics: a graph-theoretical approach[END_REF][START_REF] Tischendorf | On the usage and measurement of landscape connectivity[END_REF]. In particular, a low management intensity of the agricultural matrix (e.g., with grasslands, which are expected to be more permeable than intensively cultivated croplands) and the presence of ecological corridors (e.g., hedgerows, which are thought to facilitate dispersal of forest species among forest patches) will increase colonization rates, while decreasing local extinction rates, in many taxa, including arthropods (e.g., [START_REF] Baudry | Multi-Scale Control of Carabid Assemblages in Hedgerow Network Landscapes[END_REF][START_REF] Seibold | Arthropod decline in grasslands and forests is associated with landscape-level drivers[END_REF]. Reduced connectivity between forest patches represents a major threat to species that are forest specialists, including arthropods, and which often exhibit low dispersal abilities [START_REF] Aviron | Connectivity of cropped vs. semi-natural habitats mediates biodiversity: a case study of carabid beetles communities[END_REF]. As a result, colonization rate of forest patches may be disproportionately low compared to extinction rate and metacommunity functioning could be disrupted. In case of neutral [START_REF] Hubbell | Neutral theory in community ecology and the hypothesis of functional equivalence[END_REF] or niche differentiation processes [START_REF] Macarthur | The limiting similarity, convergence, and divergence of coexisting species[END_REF][START_REF] Maire | Habitat filtering and niche differentiation jointly explain species relative abundance within grassland communities along fertility and disturbance gradients[END_REF], such a disequilibrium can ultimately lead to an increased dissimilarity in species composition between forest patches, due to species loss or/and species replacement [START_REF] Hendrickx | Pervasive effects of dispersal limitation on within-and amongcommunity species richness in agricultural landscapes[END_REF], thereby increasing the magnitude of difference between the regional species pool and composition of local assemblages. On the contrary, in case of landscape-level filtering, species with similar trait values might be selected, leading to species homogenization at the landscape scale [START_REF] Keddy | Assembly and response rules: two goals for predictive community ecology[END_REF][START_REF] Maire | Habitat filtering and niche differentiation jointly explain species relative abundance within grassland communities along fertility and disturbance gradients[END_REF].

In addition to the degree of spatial connectivity, temporal connectivity through the age of forest patches has repeatedly been shown to influence community composition, with ancient forest patches (i.e., patches that have continuously existed for centuries) harboring a higher number of forest-specialist species with limited dispersal abilities than recent forest patches (i.e., patches that have established on former agricultural lands in the last decades) [START_REF] Dekoninck | Hoverfly communities on former agricultural fields: a study of afforestation and planted forests stands in the Voeren region[END_REF][START_REF] Desender | Diversity and assemblages of Carabid beetles in ancient forests and afforested former agricultural land[END_REF][START_REF] Debnár | Group selection harvesting supports the diversity of epigaeic arthropod assemblages (Coleoptera: Carabidae; Araneidae: Araneae; Isopoda: Oniscidae)[END_REF][START_REF] Lelli | Biodiversity response to forest structure and management: Comparing species richness, conservation relevant species and functional diversity as metrics in forest conservation[END_REF]. Ancient forest patches usually support more diverse plant assemblages than recent ones, given their higher habitat heterogeneity [START_REF] Brunet | Plant colonization in heterogeneous landscapes: an 80-year perspective on restoration of broadleaved forest vegetation[END_REF][START_REF] Burrascano | It's a long way to the top: Plant species diversity in the transition from managed to old-growth forests[END_REF] and benefit from longer duration for accumulating forest specialist species (i.e., the species-time relationship; [START_REF] Almoussawi | Forest fragmentation shapes the alpha-gamma relationship in plant diversity[END_REF][START_REF] Valdés | High ecosystem service delivery potential of small woodlands in agricultural landscapes[END_REF]. However, compared to plant species, arthropod species have received little attention with respect to species richness in ancient vs. recent forest patches [START_REF] Schowalter | Arthropod Diversity and Functional Importance in Old-Growth Forests of North America[END_REF]. Besides, to the best of our knowledge, only few studies have compared the diversity patterns of such arthropod assemblages in different contexts of landscape mosaic (e.g., [START_REF] Hendrickx | Pervasive effects of dispersal limitation on within-and amongcommunity species richness in agricultural landscapes[END_REF][START_REF] De Smedt | Linking macrodetritivore distribution to desiccation resistance in small forest fragments embedded in agricultural landscapes in Europe[END_REF].

Among major arthropod groups living in forest ecosystems, carabid beetles (Coleoptera: Carabidae) are important for ecosystem functioning and biocontrol [START_REF] Thiele | Carabid Beetles in Their Environments[END_REF][START_REF] Kromp | Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation impacts and enhancement[END_REF][START_REF] Kotze | Forty years of carabid beetle research in Europe-from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation[END_REF], and are often used as bioindicators of good forest management practices ensuring healthy and sustainable forest systems (e.g., [START_REF] Rainio | Ground beetles (Coleoptera: Carabidae) as bioindicators[END_REF][START_REF] Pearce | The use of ground beetles (Coleoptera: Carabidae) and spiders (Araneae) as bioindicators of sustainable forest management: a review[END_REF]. Forest carabid species are highly sensitive to reduction of forest patch size, as smaller patches exhibit lower habitat quality and greater edge effects than larger patches. Forest fragmentation usually induces a complete replacement of large specialist species characterized by their low mobility by small, generalist, and highly mobile species coming from surrounding habitats [START_REF] Niemelä | Carabid beetles (Coleoptera: Carabidae) and habitat fragmentation: a review[END_REF][START_REF] Rainio | Ground beetles (Coleoptera: Carabidae) as bioindicators[END_REF][START_REF] Lövei | The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands[END_REF][START_REF] Do | The effect of fragmentation and intensive management on carabid beetles in coniferous forest[END_REF][START_REF] Jung | Trait-specific response of ground beetles (Coleoptera: Carabidae) to forest fragmentation in the temperate region in Korea[END_REF]. Variations in carabid composition and diversity between mature and young forest stands have previously been shown (e.g., [START_REF] Fountain-Jones | Living near the edge: being close to mature forest increases the rate of succession in beetle communities[END_REF], as well as the interaction with forest management (e.g., [START_REF] Sroka | Ground beetle diversity in ancient woodland remnants in north-western Germany (Coleoptera, Carabidae)[END_REF][START_REF] Barkley | Does variable stand structure associated with multi-cohort forests support diversity of ground beetle (Coleoptera, Carabidae) communities in the central Nearctic boreal forest[END_REF]. Surprisingly, the role of historical continuity (i.e., forest patch age) has been largely neglected so far (but see [START_REF] Assmann | The ground beetle fauna of ancient and recent woodlands in the lowlands of northwest Germany (Coleoptera, Carabidae)[END_REF][START_REF] Desender | Diversity and assemblages of Carabid beetles in ancient forests and afforested former agricultural land[END_REF][START_REF] Hülsmann | Land-use legacy and tree age in continuous woodlands: weak effects on overall ground beetle assemblages, but strong effects on two threatened species[END_REF].

Local climatic conditions [START_REF] Park | Local climate mediates spatial and temporal variation in carabid beetle communities in three forests in Mount Odaesan, Korea[END_REF], as well as current large-scale macroclimatic gradients (e.g., [START_REF] Heino | Elements of regional beetle faunas: faunal variation and compositional breakpoints along climate, land cover and geographical gradients[END_REF] and the biogeographic history of the continent since the Pleistocene glaciations [START_REF] Calatayud | Glaciations, deciduous forests, water availability and current geographical patterns in the diversity of European Carabus species[END_REF][START_REF] Calatayud | Pleistocene climate change and the formation of regional species pools[END_REF], are widely acknowledged as important drivers of local carabid species assemblages. Together with local and landscape features, macroclimatic factors were shown to affect carabid assemblages individually, but their relative importance is still unknown, despite their crucial importance for understanding all processes underlying local arthropod, assemblages. Whether common patterns are observed across various carabid guilds remains an unanswered question. Given the importance of small deciduous forest patches as biodiversity reservoirs in agricultural landscapes [START_REF] De Blois | Landscape issues in plant ecology[END_REF] and in delivering multiple ecosystem services to human societies [START_REF] Decocq | Ecosystem services from small forest patches in agricultural landscapes[END_REF], understanding community assembly processes in these habitats embedded in patchy landscapes is particularly relevant to conservation ecology and sustainable landscape planning.

Here we aim at explaining patch-scale species richness (hereafter α-diversity) and magnitude of difference between landscape-scale species pool and patch-scale carabid assemblages (hereafter β-diversity) in small forest patches embedded in contrasted agricultural landscapes along a broad macroclimatic gradient across temperate Europe. We separated carabid assemblages into four different guilds based on habitat preferences, namely: forest specialists, forest generalists, open-habitat species, and eurytopic species. The first two guilds group "forest species", whilst the latter two group "non-forest species". More specifically, we address the following research hypotheses: (i) macroclimatic conditions act as the main ecological filter on all carabid beetle assemblages; (ii) landscape management intensity decreases local species richness (α-diversity), and increases the magnitude of difference between the landscape species pool and local assemblages (β-diversity) of forest species (particularly for specialists), whilst the reverse is true for non-forest species (particularly for open-habitat species); and (iii) patch size and patch age both positively affect α-diversity, especially for forest species, and forest specialists in particular. To test these hypotheses, we quantified the relative importance of potential environmental drivers acting at different spatial scales (patch, landscape, continent) on local carabid assemblages, using a large, dedicated field survey which encompasses seven regions distributed along a transect running from southern France to central Sweden.

METHODS

Study sites

We collected data across a total of 221 deciduous forest patches, distributed among seven European regions. Regions were distributed along a south-west-to-north-east gradient of ca.

2,100 km across the European temperate forest biome (Fig. 1). In each region, we selected two 5 km × 5 km landscape windows differing by their degree of landscape permeability (see [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF] for more details). The first window was characterized by forest patches embedded in an intensively cultivated open field landscape (hereafter "open field"). The second window contained forest patches more or less connected by woody corridors (e.g., hedgerows) within a less intensively managed landscape, dominated by grasslands and small crop fields (hereafter "bocage"). A detailed and updated list of landscape and macroclimatic variables associated with each landscape window can be found in [START_REF] Vanneste | Functional trait variation of forest understorey plant communities across Europe[END_REF].

For each landscape window, we computed area, perimeter, and age of all forest patches using digitized maps (one contemporary map at a scale of 1:25,000 and historical maps from the 17 th , 18 th , 19 th , and 20 th centuries) in a Geographic Information System (GIS; ARCGIS 9.3, ESRI).

Patches were subsequently distributed among four classes according to their area and age: small (< median patch area value for the focal window) and recent (< 150 years); large (> median patch area value for the focal window) and recent; small and ancient (> 150 years); and large and ancient.

# Figure 1 here #

Carabid sampling and habitat preference

Whenever possible, we selected four forest patches (i.e., four repetitions) per level of patch sizeage combination (n = 4) and per landscape window (n = 14). A perfect balanced design was achieved in five out of seven regions, to finally include a total of 221 forest patches to trap carabid beetles (i.e., 16 forest patches in each window, except 14 in the openfield window in eastern Germany and 15 in the openfield window in southern Sweden). For this purpose, we used 10-cm diameter pitfall traps installed for 14 consecutive days in both spring (ca. April) and summer (ca. August) 2013. Traps were filled with 200 mL of a 50 % conservative solution of ethylene-glycol and a few drops of detergent, and protected from litter and rain fall by aluminum roofs. A total of four pairs of traps were disposed in each patch as follows. A first pair, consisting of two traps separated by a plastic barrier (100-cm long, 18-cm high), was disposed into the inner part of a south-facing edge (or, when not possible, first a west-, and then an east-facing edge was chosen). This setup was replicated 5-m apart along the same edge. A third pair was installed at the barycenter of the forest patch (except in eastern Germany, where all traps were located in the edge), and similarly replicated (fourth pair). The plastic barrier was always parallel to the selected forest edge. To make data comparable among the seven studied regions, and because of the latitudinal climatic gradient covered by our study, the two sampling sessions carried out in each region started when local growing degree hours (GDH ; [START_REF] Graae | On the use of weather data in ecological studies along altitudinal and latitudinal gradients[END_REF] reached values of ca. 10,000°C h and 20,000°C h, respectively. Following trap collection, carabid beetles were sorted in the lab in a 70% ethanol solution and identified to the species level following [START_REF] Jeannel | Coléoptères Carabiques, Faune de France[END_REF][START_REF] Jeannel | Coléoptères Carabiques II, Faune de France[END_REF]. Species names follow Fauna Europaea [START_REF] De Jong | Fauna Europaea-all European animal species on the web[END_REF]. Data from all pitfall traps of a given patch and from the two trapping sessions were pooled in all subsequent statistical analyses.

Species were distributed among four guilds, according to their habitat preference and using knowledge from the scientific literature [START_REF] Hůrka | Carabidae of the Czech and Slovak Republics, Vit Kabourek Editors[END_REF][START_REF] Sadler | Investigating environmental variation and landscape characteristics of an urban-rural gradient using woodland carabid assemblages[END_REF][START_REF] Gaublomme | The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape[END_REF][START_REF] Bräunicke | Lebensraumpräferenzen der Laufkäfer Deutschlands-Wissensbasierter Katalog[END_REF]: forest-specialist species, limited to stable, mature forest stands; forest-generalist species, occurring in any type of forest stand, in ancient as well as recent forest; open-habitat species, associated with non-forest habitats such as grasslands and arable lands; and eurytopic species, occurring in open habitats and tolerating transiently forest habitats.

We could not assign only one species (Oodes helopioides, n = 1 individual) to any group because of a lack of information in the literature.

Environmental variables

Three groups of explanatory variables (patch, landscape, and macroclimatic variables) were derived from field observations, historical archives, or global climatic layers, for and around each forest patch using a GIS.

Macroclimatic variables

To assess the influence of macroclimate on species diversity, we extracted 10 candidate bioclimatic variables from the WorldClim global database (1-km resolution, http://www.worldclim.org), and averaged each variable for each forest patch using all 1-km 2 pixels intersecting it. Four macroclimatic variables were retained for further analyses, based on a principal component analysis (PCA; see [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF], namely maximum temperature of the warmest month (MaTWm; BIO5); minimum temperature of the coldest month (MiTCm; BIO6); precipitation of the wettest month (PWm; BIO13); and precipitation of the driest month (PDm; BIO14). The selection was made in a way to minimize the correlation between variables and to maximize the correlation with the PCA axes. Our variables were correlated (Pearson"s r) as follow with the PC1 (67.0 % of explained variance) and PC2 (17.2 %): BIO5 (PC1: r = -0.64; PC2: r = -0.12), BIO6 (PC1: -0.93; PC2: 0.34), BIO13 (PC1: -0.52; PC2: -0.85), and BIO14 (PC1: -0.94; PC2: -0.04).

Landscape variables

Landscape variables were computed for five concentric "doughnut"-like buffers of increasing width around each forest patch: 50; 100; 250; 500; and 1,000 m width. We used Corine Land Cover 2006 [START_REF] Büttner | CLC2006 technical guidelines[END_REF] to map the distribution of woodland, cropland, and grassland. We digitized hedgerows from aerial photographs. As proposed by [START_REF] Martin | Measuring and selecting scales of effect for landscape predictors in species-habitat models[END_REF] and [START_REF] Fahrig | Rethinking patch size and isolation effects: the habitat amount hypothesis[END_REF], we considered composition-based measurements of spatial isolation for each forest patch, by calculating the proportion of each cover type and the hedgerow density within each buffer.

Patch-scale attributes

We included patch area and patch age as covariates in our analyses, to account for both the species-area and species-time relationship [START_REF] Rosenzweig | Species diversity in space and time[END_REF]. We took the coefficient of variation in elevation values (CVe) within a given forest patch using the ASTER Global Digital Elevation Map at 30-m resolution (see [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF] for further details), as a proxy for heterogeneity of abiotic conditions (incl. microclimate, soil conditions, and light availability) [START_REF] Lenoir | Climatic microrefugia under anthropogenic climate change: implications for species redistribution[END_REF][START_REF] Graae | Stay or go-how topographic complexity influences alpine plant population and community responses to climate change[END_REF]. Finally, we computed a dissimilarity index in understory plant species composition within each forest patch (i.e., intra-patch -diversity; see [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF] for details on computation), separately for forest plant specialists and generalists following distinction criteria as in [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF].

Data analysis

All statistical analyses were performed in R 3.5.0 (R. [START_REF] Team | R: A language and environment for statistical computing[END_REF].

Multiple-site dissimilarities

To evaluate whether between-patches differences in carabid composition were due to replacement of species or to nestedness (since the same sampling effort was applied irrespective to patch size), we computed multiple-site dissimilarities, separating the turnover and nestednessresultant components of overall Sørensen-based multiple-site dissimilarity as proposed by [START_REF] Baselga | Partitioning the turnover and nestedness components of beta diversity[END_REF][START_REF] Baselga | The relationship between species replacement, dissimilarity derived from nestedness, and nestedness[END_REF]. Dissimilarities were evaluated at two different scales: (1) between-patch dissimilarity within each landscape window, and (2) between-region dissimilarity along the European gradient. We used the "betapart" package [START_REF] Baselga | betapart: Partitioning beta diversity into turnover and nestedness components[END_REF].

Species diversity

We computed patch-level diversity, defined as the total number of species trapped per forest patch; landscape-level diversity, defined as the total number of species retrieved in all patches of a given landscape window; and diversity, defined as the magnitude of difference between the landscape-level species pool (diversity) and the composition of the local assemblage (diversity): (. Diversity values were calculated separately for each of the four carabid guilds, using the "vegan" package [START_REF] Oksanen | Community Ecology Package. R package version 2[END_REF].

Model selection. The effects of patch, landscape, and macroclimatic attributes on diversity and diversity were quantified using generalized linear mixed-effects models (GLMMs) and the "lme4" package [START_REF] Bates | Fitting Linear Mixed-Effects Models Using lme4[END_REF]. Negative binomial distribution models (including an observation level factor to account for overdispersion) were used for patch-level diversity.

Gaussian error distributions with an identity link were used for diversity. Distribution families were chosen using the functions descdist and fitdist of the "fitdistrplus" package [START_REF] Delignette-Muller | fitdistrplus: An R package for fitting distributions[END_REF]. To address possible multi-collinearity issues between macroclimatic, landscape, and patch attributes, we computed pairwise Pearson correlation tests between all continuous variables, and considered a threshold of 0.70 to consider two independent predictors as correlated [START_REF] Dormann | Collinearity: a review of methods to deal with it and a simulation study evaluating their performance[END_REF]Appendix S1). All explanatory variables were standardized prior to analyses by subtracting the mean and dividing by the standard deviation. We included "region" and "window type" (nested within "region") as random intercept terms in all models to account for the hierarchical structure of the sampling design and potential spatial autocorrelation between assemblages occurring in the same region and landscape window.

In a first step, we selected the most relevant spatial scale (from 50 to 1,000 m) to consider for each landscape variable (i.e., hedgerow density, grassland, crop, and forest relative proportion) at each diversity level ( and diversity) and for each guild. For each landscape variable, we ran a model at each spatial scale (one variable at a time) and retained the one scale from the model with the lowest Akaike Information Criterion (AIC; [START_REF] Burnham | Multimodel inference: understanding AIC and BIC in model selection[END_REF].

In a second step, we constructed complete models regrouping all explanatory variables (see "Environmental variables"): first order (BIO X ) and second order (BIO X 2 ; to allow for non-linear macroclimatic variation) macroclimatic variables (n = 8); landscape variables (n = 4); and patch variables (n = 5). Because of the large number of variables, we could not compare candidate models with all possible combinations of variables. We thus performed a backward stepwise selection of fixed explanatory variables: the full model was simplified step-by-step by removing the most non-significant explanatory variables based on likelihood-ratio tests. The ML estimation was used during model selection, and then REML was used for fitting the final model.

Variation partitioning

We ran a variation partitioning procedure including all response variables to quantify the respective explanatory power of the three groups of explanatory variables (patch, n = 5 variables; landscape, n = 4; and macroclimatic, n = 8) according to [START_REF] Legendre | Numerical ecology[END_REF] and [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF]. We constructed GLMMs (diversity) or LMMs (diversity) including combinations of one, two, or three variable groups plus the random effects, and determined the proportion of variation explained by the fixed variables (marginal R 2 ; R 2 m ) according to [START_REF] Nakagawa | A general and simple method for obtaining R2 from generalized linear mixed-effects models[END_REF]. We quantified the proportion of this variation explained by the unique and shared contribution of patch, landscape, and macroclimatic variables, and expressed this relative to the total amount of variation explained by the fixed factors in the global model (containing all three groups of explanatory variables).

RESULTS

A We observed important dissimilarities in species composition among regions along the European gradient, irrespective of the guild considered (Sørensen dissimilarity: 0.62-0.72). These were mainly driven by species turnover (Simpson dissimilarity: 0.50-0.61) (Fig. 2). Similar results

were found among patches within each landscape window and for each guild, except for forestspecialist species, for which the nestedness component of Sørensen dissimilarity almost equaled the turnover component (Fig. 2).

# Figure 2 here #

Species richness (α-diversity)

Species richness varied along the latitudinal gradient for all guilds, with a trend toward more species at intermediate than at extreme latitudes along our European gradient (Fig. 3). In most regions, open-habitat species richness was (or tended to be) higher in open field than in bocage windows (Fig. 3d). We found no significant difference in mean species richness of the three other guilds between open field and bocage windows, irrespective of the region considered (except in northern France: less eurytopic species in bocage than open field window; Fig. 3c).

# Figure 3 here # Based on the marginal and conditional R 2 difference derived from mixed-effects models, and contrary to other guilds, variance in forest-specialist species richness was mostly explained by random factors (i.e., region and window type nested in region) (Fig. 4a-d; Table 1a). Among all groups of explanatory variables, the pure effect of macroclimate explained the largest proportion of local species richness (α-diversity) for all guilds (89.8%, 84.3%, and 75.5% of the explained variance for forest-specialist, eurytopic, and open-habitat species, within the fixed part of the model, respectively; Fig. 4a,c,d) except forest generalists. The percentage of variation explained by patch-scale attributes and landscape variables was negligible. Variation in forest-generalist species richness was better explained by the pure (50.3%) and total (86.5%) effect of patch-scale attributes, than by pure (19.0%) and total (49.2%) effects of macroclimatic variables (Fig. 4b).

# Figure 4

Magnitude of difference between local assemblages and the landscape pool (β-diversity)

Overall, we found high values of β-diversity among fragments, irrespective of guild, landscape type and region considered (Fig. 5). We found the lowest magnitude of difference between the landscape pool and local assemblages for forest-specialists, especially at intermediate latitudes (Fig. 5a).

# Figure 5 here #

Our model explained a low proportion of the observed variation in β-diversity of carabid species assemblages for all guilds and especially for open-habitat species (R 2 m = 0.096; R 2 c = 0.232) (Fig. 4e-h; Table 1b).

For all guilds, β-diversity was mostly explained by the pure effect of macroclimatic variables (from 31.9% to 71.7% of the explained variance; Fig. 3e-h), except for open-habitat species (pure effect of patch-sale attributes: 41.2%). For all guilds except forest-specialist species, landscape variables had a non-negligible effect on β-diversity (pure effect from 13.4% to 22.9%; Fig. 4f-h).

The magnitude of difference between the landscape pool and local assemblages was primarily explained by extreme temperature (forest-specialist, generalist, and eurytopic species) and extreme rainfall (forest-specialist, eurytopic, and open-habitat species) conditions (Table 1b).

Beta-diversity of forest-generalist species decreased with increasing maximum temperature during the warmest month (MaTWm), whilst the reverse pattern was observed for eurytopic species. Both forest-specialist and forest-generalist β-diversity increased with minimum temperature during the coldest month (MiTCm). Beta-diversity of eurytopic species peaked in regions with intermediate amount of precipitation during the wettest month (PWm), and increased with the amount of precipitation during the driest month (PDm). The latter also had a positive effect on forest-specialist β-diversity.

Landscape characteristics affected β-diversity of all guilds except forest-specialist species (Table 1a). Beta-diversity of forest-generalist species increased with hedgerow density (50 m) and the proportion of forest (500 m) in the landscape. Beta-diversity of eurytopic species decreased with increasing proportion of crops (250 m) and grassland (1,000 m) in the surrounding landscape. Beta-diversity of open-habitat species decreased with the increasing amount of crops (500 m) and forest (50 m) in the surrounding landscape.

Beta-diversity of forest specialists and forest generalists was higher in recent than ancient forest patches (Table 1a). Topographic variability (CVe) had a negative effect on forest-specialist β-diversity, but a positive effect on eurytopic and open-habitat β-diversity. Beta-diversity of forest-generalist species increased with β-diversity of generalist plant species in the herb layer but decreased with increasing β-diversity of forest specialist plants. The opposite pattern was found for open-habitat β-diversity.

DISCUSSION

In this study, we have quantified the respective importance of forest patch characteristics, landscape features, and macroclimate in driving αand β-diversities of carabid beetle assemblages. Both within and between small forest patches embedded in contrasted agricultural landscapes. We showed that both diversity components were primarily determined by macroclimate, likely via the size and composition of regional species pools. Local carabid species assemblages differed because of habitat preferences. While the diversity of non-forest species (open-habitat and eurytopic) in forest patches was mostly influenced by landscape features, especially proportion of different land-cover types, the diversity of forest species (specialist and generalist) was on the other hand positively affected by patch-scale attributes, such as patch age and heterogeneity in patch properties (abiotic and biotic).

Macroclimate: a prominent ecological filter

We found support for our first research hypothesis: macroclimatic conditions act as the main ecological filter on carabid beetle diversity patterns, as implied by the high relative importance values of macroclimatic factors in explaining local αand -diversity patterns. More precisely, extremes of temperature (cf. MaTWm, MiTCm) and, to a lesser degree, extremes of precipitation (cf. PWm, PDm), emerge as crucial drivers of carabid diversity. This is consistent with earlier studies, showing that the strong effect of latitude on carabid species richness is mediated by climate, and then more by temperature than precipitation (e.g., [START_REF] Schuldt | Environmental and historical effects on richness and endemism patterns of carabid beetles in the western Palaearctic[END_REF][START_REF] Ernst | Drivers and patterns of ground-dwelling beetle biodiversity across Northern Canada[END_REF]. In addition, and congruently, regional species pools of all species guilds tend to show a hump-shaped relationship with lower regional species richness at both extremes of the latitudinal gradient (Appendix S3), while a high species turnover is observed among regions along this gradient (Fig. 2). Furthermore, distinct carabid species respond differently to macroclimatic factors, depending on habitat preference.

Forest species mostly respond to annual extreme temperatures, with the α-diversity of specialist and generalist species increasing as MiTCm and MaTWm decreased and increased, respectively. Cold temperatures (cf. MiTCm) are known to exert strong physiological constraints on overwintering poikilothermous species [START_REF] Thiele | Carabid Beetles in Their Environments[END_REF][START_REF] Lövei | Ecology and behavior of ground beetles (Coleoptera: Carabidae)[END_REF]. The majority of temperate carabid species require winter diapause to complete larval development or gonad maturation and cannot survive environments with too mild winter temperatures, since diapause is not triggered [START_REF] Kotze | Forty years of carabid beetle research in Europe-from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation[END_REF]. On the other hand, cold induces physiological costs and only few species have adapted mechanisms to survive very harsh winter conditions [START_REF] Bale | Insect cold hardiness: a matter of life and death[END_REF]. High temperature (cf. MaTWm) usually increases activity and development rates (e.g., [START_REF] Taylor | Analysis of the effect of temperature on insects in flight[END_REF][START_REF] Forrest | Complex responses of insect phenology to climate change[END_REF] and throughout the active period of the year, insects that face warm conditions during their ontogenetic development can acclimate to even higher temperatures [START_REF] Sheikh | Diverse adaptations in insects: A Review[END_REF]. Warmer weather conditions may thus promote species movements within landscapes, thereby affecting β-diversity patterns (here defined as the magnitude of difference between the landscape-level species pool and the composition of local assemblages).

Among non-forest species, eurytopic species respond to both temperature (cf. negative effect of both MaTWm and MiTCm on α-diversity) and precipitation (cf. negative effect of both PWm and PDm on α-diversity), while open-habitat α-diversity rather show a unimodal relationship with MiTCm and PDm. Eurytopic species can live and disperse in most habitats, but may not have developed adaptations for extreme climatic conditions. Moreover, as non-forest species, eurytopic and open-habitat species cannot benefit from canopy-induced microclimatic conditions of the understory, which is typically moister over the year, cooler in summer, and milder in winter [START_REF] De Frenne | Global buffering of temperatures under forest canopies[END_REF]. When the temperature is too high or the environment too dry, these species are thus more prone to enter into summer diapause or quiescence to face unsuitable conditions [START_REF] Masaki | Summer diapause[END_REF]. Because of reduced activity, these species likely disperse less efficiently, thereby increasing compositional dissimilarity among patches.

As a consequence of the current climate change and because of physiological constraints of species to their regional climate, the elevational range of some carabid species has changed in the past 30 years, to track the shifting isotherms (e.g., [START_REF] Pizzolotto | Changes in ground beetle assemblages above and below the treeline of the D olomites after almost 30 years (1980/2009)[END_REF][START_REF] Moret | Climate warming effects in the tropical Andes: first evidence for upslope shifts of Carabidae (Coleoptera) in Ecuador[END_REF].

Movement of species northward (in the Northern Hemisphere) can also be expected, but changes in latitudinal distributions can be hard to predict as the dominant diapause-inducing cue (photoperiod) will not be affected by climate change while other cues (temperature, moisture)

will [START_REF] Huffeldt | Photic Barriers to Poleward Range-shifts[END_REF]. Species may indeed suffer from mismatched information between critical photoperiod for diapause induction and other environmental conditions acting on the physiology, development, or behavior of specific life stages, which could affect their survival [START_REF] Bale | Insect overwintering in a changing climate[END_REF][START_REF] Tougeron | How climate change affects the seasonal ecology of insect parasitoids[END_REF]. In addition, by causing species-specific shifts in phenology, climate change can alter interspecific interactions as well [START_REF] Damien | Prey-predator phenological mismatch under climate change[END_REF].

The legacy of the biogeographical history of Europe (particularly glaciations of the Pleistocene and the subsequent post-glacial recolonization of northern regions) may confound the pure effect of current climatic conditions in explaining the macroecological diversity patterns we observed. This is particularly true for dispersal-limited organisms such as carabid beetles [START_REF] Calatayud | Glaciations, deciduous forests, water availability and current geographical patterns in the diversity of European Carabus species[END_REF][START_REF] Calatayud | Pleistocene climate change and the formation of regional species pools[END_REF], and may explain the lower species richness recorded in the northern regions.

Landscape features explain the contribution of non-forest species to local assemblages

Our results reveal that the composition of the landscape matrix into which forest patches are embedded significantly explains the αand -diversity of non-forest carabid species (cf.

eurytopic and open-habitat guilds). In particular, focal patches surrounded by forest and grassland have a negative effect on species richness for these species. Consistently, the number of openhabitat species was higher in open field landscapes compared to bocage landscapes. This may be explained by the diversity of crops cultivated in the surrounding fields (e.g., cereals, sugar beet, potato, rapeseed), each associated with a particular pool of species [START_REF] Holland | Farm-scale spatiotemporal dynamics of predatory beetles in arable crops[END_REF][START_REF] Marrec | Crop succession and habitat preferences drive the distribution and abundance of carabid beetles in an agricultural landscape[END_REF], whilst grasslands may harbor less specific carabid assemblages and benefit from spillover from adjacent habitats, including forests [START_REF] Schneider | Spillover from adjacent crop and forest habitats shapes carabid beetle assemblages in fragmented semi-natural grasslands[END_REF][START_REF] Duflot | Adjacent woodlands rather than habitat connectivity influence grassland plant, carabid and bird assemblages in farmland landscapes[END_REF].

Moreover, arable lands may allow beetles to move more easily on the ground than grasslands, particularly because of differences in vegetation density [START_REF] Thomas | Carabid movement and vegetation density: Implications for interpreting pitfall trap data from split-field trials[END_REF]. Our results are consistent with previous studies, which showed that landscape attributes affect biodiversity in important ways (see [START_REF] Fletcher | Divergent perspectives on landscape connectivity reveal consistent effects from genes to communities[END_REF] for a review). Open-habitat and eurytopic species use forest habitats as alternative, temporary habitat, but are less competitive there than in more open habitats (e.g., [START_REF] Niemelä | Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae) in western Canada[END_REF]. In particular, they enter forest edges to find a shelter, to search for food or to seek overwintering sites [START_REF] Knapp | Temporal changes in the spatial distribution of carabid beetles around arable field-woodlot boundaries[END_REF]. The decreased magnitude of difference between the landscape-level species pool and the composition of local assemblages with increasing crop or/and grassland cover in the landscape suggests that these habitats allow a large suite of non-forest species to enter forest patches. However, the high turnover observed among patches of a given landscape window suggests that they are not present at the same time in most patches.

With the exception of -diversity of forest-generalist species, which increases with hedgerow density and forest cover in the surrounding landscape, landscape factors do not affect the αand -diversity patterns of forest-specialist and generalist carabid species. This result indicates that hedgerows are not always positive for species movement (e.g., [START_REF] Baudry | Multi-Scale Control of Carabid Assemblages in Hedgerow Network Landscapes[END_REF].

Alternatively, hedgerows in the studied landscapes may be too recent or scarce to serve as highquality corridors [START_REF] Closset-Kopp | Using process-based indicator species to evaluate ecological corridors in fragmented landscapes[END_REF][START_REF] Lenoir | Historical continuity and spatial connectivity ensure hedgerows are effective corridors for forest plants: evidence from the species-time-area relationship[END_REF] for forest carabid species, that are well known for their very low dispersal capacities [START_REF] Rainio | Ground beetles (Coleoptera: Carabidae) as bioindicators[END_REF]. However, hedgerows can allow some forest species to disperse between forest patches as previously suggested (e.g., [START_REF] Baudry | Multi-Scale Control of Carabid Assemblages in Hedgerow Network Landscapes[END_REF], increasing the probability that more species will reach forest patches. However, the high nestedness we found for forest-specialist species compared to other guilds suggests that only a few species are actually able to colonize new forest patches.

Increased magnitude of difference between the landscape-level species pool and the composition of local assemblages in bocage landscapes could then be linked to random local species replacement due to competition, or neutral processes. In summary, we found little support to our second research hypothesis: landscape management intensity only weakly influences the diversity of forest carabid species, but instead increases the number of non-forest species in small forest patches.

Forest patch characteristics drive forest carabid species diversity

We found that patch-scale attributes mostly influence the αand -diversity patterns of forest carabid species. Patch characteristics were even the primary drivers of α-diversity for forestgeneralist species. This includes patch age and various proxies for local environmental heterogeneity. The higher number of forest generalists in ancient compared to recent forest patches likely reflects the accumulation of weaker dispersers over time, according to the speciestime relationship [START_REF] Rosenzweig | Species diversity in space and time[END_REF], which has already been showed for vascular plant species in the same study system [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF]. Similarly, forest carabid species are large species, with poor dispersal abilities, and long life cycles [START_REF] Thiele | Carabid Beetles in Their Environments[END_REF][START_REF] Rainio | Ground beetles (Coleoptera: Carabidae) as bioindicators[END_REF].

Ancient forests are thought to be more stable, allowing these species to persist [START_REF] Driscoll | Beetle responses to habitat fragmentation depend on ecological traits, habitat condition, and remnant size[END_REF][START_REF] Schowalter | Arthropod Diversity and Functional Importance in Old-Growth Forests of North America[END_REF]. Moreover, ancient forest patches are also thought to exhibit more heterogeneous environmental conditions than recent patches [START_REF] Lawesson | Species diversity and area-relationships in Danish beech forests[END_REF][START_REF] Honnay | Effects of area, age and diversity of forest patches in Belgium on plant species richness, and implications for conservation and reforestation[END_REF][START_REF] Schowalter | Arthropod Diversity and Functional Importance in Old-Growth Forests of North America[END_REF], thereby providing carabid beetles with more potential niches. This is confirmed by the positive effect of plant diversity (cf. intra-patch -diversity of forest specialists) on the species richness of forest generalist carabid beetles, since plant species richness has also been shown to increase with patch heterogeneity [START_REF] Jamoneau | Drivers of plant species assemblages in forest patches among contrasted dynamic agricultural landscapes[END_REF].

Carabid beetles are proven to be very sensitive to variations in forest characteristics, so that their assemblages change during the forestry cycle, according to variation in forest structure and composition (e.g., [START_REF] Butterfield | Carabid community succession during the forestry cycle in conifer plantations[END_REF][START_REF] Koivula | Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient[END_REF][START_REF] Magura | Diversity and composition of carabids during a forestry cycle[END_REF].

The -diversity of forest generalists was influenced by the same patch-scale factors as diversity, but in the opposite direction. This indicates that the few forest-generalist species present in a given landscape window tend to occupy all ancient, heterogeneous forest patches of this window. Interestingly, the -diversity of forest specialists also decreased as patch age increased, consistent with species accumulation over time. However, local species richness was mostly determined by region and landscape-window type (i.e., random factors in our models).

This result suggests that the colonization of forest patches within landscape windows occurs at random in such fragmented systems where metapopulation dynamics for these dispersal-limited species are likely disrupted. The positive effect of local topography (another proxy for intra-patch heterogeneity) on local species richness can be explained by the fact that small-scale variation in topography offers more thermal variability than flat areas [START_REF] Lenoir | Local temperatures inferred from plant communities suggest strong spatial buffering of climate warming across N orthern E urope[END_REF], as well as more micro-habitats (e.g. various depth and quality of litter, coarse woody debris, snags, different soil texture; [START_REF] Dwyer | Influence of topographic heterogeneity on deciduous litter decomposition[END_REF]. These features have already been shown to affect carabid assemblages [START_REF] Latty | Response of ground beetle (Carabidae) assemblages to logging history in northern hardwood-hemlock forests[END_REF][START_REF] Tyler | Differences in abundance, species richness, and body size of ground beetles (Coleoptera: Carabidae) between beech (Fagus sylvatica L.) forests on Podzol and Cambisol[END_REF], and other arthropods such as ticks, Ixodes ricinus [START_REF] Ehrmann | Environmental drivers of Ixodes ricinus abundance in forest fragments of rural European landscapes[END_REF].

It is noteworthy that patch area was not significant in our models, which is likely a consequence of our sampling design, since we applied the same sampling effort irrespective of patch size, using traps that can catch beetles over a very limited area relative to patch size.

Nonetheless, recent reviews suggested that small and isolated habitat patches can have similar ecological values as large patches, especially when they encompass the same overall area at the landscape to regional scale [START_REF] Fahrig | Ecological responses to habitat fragmentation per se[END_REF][START_REF] Wintle | Global synthesis of conservation studies reveals the importance of small habitat patches for biodiversity[END_REF].

To a lesser extent, patch-scale attributes also impact the diversity of open-habitat species. In particular, the magnitude of difference between the landscape-level species pool and the composition of their local assemblages was positively affected by local topography and diversity of forest plant specialists. This suggests that open-habitat species, and, to a lesser degree, eurytopic species (see positive relationship between -diversity of eurytopic species and CVe) hardly colonize heterogeneous forests established on complex terrains. Consistently, the number of open-habitat species (-diversity) decreased with increasing vegetation heterogeneity (cf. -diversity of forest specialist plants) and increasing elevational heterogeneity (cf. CVe).

Based on these results, we can partly confirm our third research hypothesis: patch age, not patch size (at least when a similar sampling effort is applied irrespective of patch size), increases α-diversity of forest species, but local abiotic and biotic heterogeneities have an even more positive effect.

CONCLUSION

We show that composition of carabid assemblages in small forest patches is strongly driven by Species assembly was also importantly altered by patch environmental heterogeneity and age, as previously observed in the same forest networks for other arthropods and plants [START_REF] Valdés | The contribution of patch-scale conditions is greater than that of macroclimate in explaining local plant diversity in fragmented forests across E urope[END_REF][START_REF] De Smedt | Linking macrodetritivore distribution to desiccation resistance in small forest fragments embedded in agricultural landscapes in Europe[END_REF]). Yet, we did not find any evidence on an influence of forest size.

These results highlight the major need of protecting old forest patches embedded in agricultural landscapes, even the small ones. Finally, landscape management intensity weakly influences the diversity of forest carabid species, but increases the number of non-forest species in small forest patches, while decreasing their turnover among patches. These results invalidate our previous assumptions and instead made us consider that observed patterns, at multiple spatial scales, are likely linked to functional traits, more than species habitat preference only. Congruently, Le [START_REF] Provost | Land-use history impacts functional diversity across multiple trophic groups[END_REF] recently showed that mobility, resource-acquisition, and body-size traits drive local species assembly through environmental filtering acting at the landscape and patch levels, especially in landscapes suffering high short and long-term turnover such as agricultural landscapes. Such evidence has not been provided yet on assemblages living in small forest patches embedded in these agricultural landscapes. α-diversity) or linear mixed-effect models (LMM; β-diversity) with the predictor variables as fixed effects and "region" as well as "window type" (nested within "region") as random intercept terms.

  total of 35,072 individuals corresponding to 159 species were collected from the 221 forest patches. Seven highly frequent species were retrieved in more than 100 patches (Abax parallelepipedus (n = 2,443 individuals), Carabus hortensis (n = 968), C. nemoralis (n =1,206), Nebria brevicollis (n = 1,605), Pterostichus melanarius (n = 6,583), P. niger (n = 2,095), and P. oblongopunctatus (n = 2,926)). Ten species (n = 5,270 individuals) were forest specialist (FS), 24 species (n = 12,350) forest generalist (FG), 51 species (n = 14,803) eurytopic (EU), and 72 species (n = 2,613) open-habitat (OH) (see Appendix S2 for the full list of species together with their associated habitat-preference guild and total abundance).

  macroclimatic conditions (and possibly by confounding historical factors) for all habitatpreference guilds. This result indicates the importance of considering macroecological factors as the main drivers of local carabid species assemblages. This novel result was yet unknown for most arthropods including carabid beetles, and much needed to improve our knowledge in arthropod macroecological patterns. Indeed, in the current climate change context, integrating climatic information in large-scale (both temporal and spatial) ecology studies is required. If species facing climate change are not sufficiently plastic or adaptable in their climatic tolerance range, they would have to move northward (in the Northern Hemisphere) or upward in mountainous systems to track the shifting isotherms and survive, and consequently impact local species assemblages.

Figure captions Figure 1

 captions1 Figure captions

Figure 2

 2 Figure2Among-region along the European gradient ("All gradient") and among-patch within each region multiple-site dissimilarities. Turnover (blue; Simpson dissimilarity) and nestedness components (yellow; nestedness-resultant fraction of Sørensen dissimilarity) are distinguished.

Figure 3

 3 Figure 3 Mean (± SD) carabid species richness (α-diversity) sampled in each forest patch, grouped based on region and window type (orange: "bocage"; blue: open-field). Statistical differences between each pair of groups were evaluated using an analysis of variance model followed by a Tukey post-hoc test. FS: forest specialists; FG: forest generalists; EU: eurytopic species; OH: open-habitat species.

Figure 4

 4 Figure 4 Results of the variation partitioning for each combination of diversity levels and habitat-preference groups as the response variable. The amount of variation explained by pure and shared contributions of each variable group was calculated as a percentage of the total variation explained by the fixed factors in the global model (including the three groups of explanatory variables). Values of marginal R 2 (R 2 m ) and conditional R 2 (R 2 c ) of the global models are shown for each response variable. Variation partitioning was based on generalized (GLMM;

Figure 5

 5 Figure 5 Mean (± SD) between-patch carabid species dissimilarity (β-diversity), grouped based on region and window type (black: "bocage"; gray: open-field). Statistical differences between each pair of groups were evaluated using an analysis of variance model followed by a Tukey post-hoc test. FS: forest specialists; FG: forest generalists; EU: eurytopic species; OH: openhabitat species.

ACKNOWLEDGEMENTS

We thank Catherine Bataillon, Kent Hansson, Laurent Raison, Sabinea Sigfridsson, and Ilka Strubelt for their help during field work, Willem Proesmans, Laurent Raison, and Rieneke Vanhulle for their help for species sorting and identification. This research was funded by the ERA-Net BiodivERsA project smallFOREST, with the national funders ANR (France), MINECO (Spain), FORMAS (Sweden), ETAG (Estonia), DFG/ DLR (Germany) and

DATA AVAILABILITY STATEMENT

Full data are available in the Dryad data repository (https://doi.org/10.5061/dryad.g79cnp5ns) while total abundances per species and species habitat preferences are listed in Appendix S2.

SUPPORTING INFORMATION

Additional supporting information may be found in the online version of this article at the publisher"s website.

Appendix S1

Correlation table between all patch, landscape, and macroclimatic-level variables.

Appendix S2 Complete list of species that were found in the sampled forest patches, along with their habitat preference.

Appendix S3 Gamma-diversity of all species together and per habitat-preference guild.


est destinée au dépôt

Introduction

Michela Russo

To cite this version:

The Emergence of Grammars. A closer look at dialects between phonology and morphosyntax, Michela Russo (ed.) This book brings together ten original contributions to the linguistic study of grammars. In several works carried out according to different theoretical perspectives, linguists have tried to identify the phonological, morphological, syntactic, and semantic properties of a grammar. What is a grammar? What types of grammar are possible in natural languages? Why and to what extent do grammatical properties vary from one language to another? The answer to these questions is not obvious.

Linguistic analysis can have for object the linguistic knowledge, the grammatical competence (i.e., I-language), while an explicit grammar must model this knowledge in its entirety. A mentalist interpretation of grammar implies that it is not enough to propose any axiomatized system compatible with a given corpus.

The book also raises the question of a modular grammar. Some researchers admit in the grammar only one productive component, others postulate a modular grammar composed by several autonomous generative systems interacting with one another.

The book recalls one of the major empirical challenges to which a general theory of language structure must satisfy: to describe in detail both the variation and the absence of variation between languages, and to explain why this variation has the properties that we observe.

This book also raises the question of the status of linearity in grammar, that is to say, first of all, the question of its multiplicity or its uniqueness. The topic of linearity represents what can be called the maximum impact of phonology in grammar.

The ten contributions that make up this book were conducted independently by linguists who, without necessarily sharing the same theoretical framework, share a universalist conception of grammar, and the ambition to go beyond the simple facts in order to propose an explanation.

All languages are human and the properties of Lithuanian, Gothic, Sanskrit, Nakanai, Bosnian, Croatian and Serbian, Finnic languages, Atlantic Languages, Proto-Western Arabic and Maltese, Occitan, medieval and modern Italo-Romance grammars are therefore relevant to English or French specialists as well.

Each article of this book aims to describe and understand complex data from empirical materials collected in a language, a group of languages or a type of language.

Daniel Petit studies in his Chapter 1 ("The secondary locative cases in the Lithuanian dialect of Zietela") the Lithuanian dialect of Zietela (Belorussia), where the Common Baltic system of secondary locative cases has been maintenaid. He provides information on this variety in the context of Baltic languages, then he addresses the case system in detail. He presents the reconfiguration -thus the emergence of a new system in Zietela -of case markers in the language. This system is structured in Old Lithuanian according to two parameters [±CONTACT], [±MOTION]. In this chapter he aims to formally describe the new features configuration and distribution in the dialect of Zietela from the historical development. This dialect has original features, some of which are contact-induced innovations.

Nadira Aljović in her Chapter 2 ("First conjunct agreement is not an elliptical illusion: a case of prenominal adjectives") addresses the question of first-conjunct agreement with prenominal adjectives in Bosnian, Croatian, Serbian. She argues against an analysis of this phenomenon based on ellipsis. The claim is that first conjunct agreement can arise within a structure in which the prenominal element scopes semantically and syntactically over a conjunction noun phrase.

Vincent Surrel's Chapter 3 ("Null Subjects and subjects pronouns in diachrony: evidence from textual sources of Northern Occitan Velay") focuses on the little-known pronominal systems present in northern Occitan, intermediate between the Oc and Oïl domains.

One would expect verbs in Occitan, as in other Romance languages such as Italian or Spanish, to have no subject. The forms analysed by Vincent Surrel are therefore surprising and require a linguistic and dialectological interpretation. In the dialects of Oïl, subject pronouns are obligatory for inflected verbs, and this grammatical distinction, known by linguists as the null subject parameter (or pro-drop parameter), groups together two typological areas with or without an expressed subject. Vincent Surrel highlights the behavior of transitional dialects, with partial pro-drop. The idea is therefore to define these partial paradigms which seem to escape the traditional pro-drop parameterization. He proposes to model this pronominal variation.

Michela Russo and Shanti Ulfsbjorninn in their Chapter 4 ("Accounting for the definite articles in Medieval Italian and Modern Dialects. allomorphy -a commun UR") explore the properties of definite articles in Italo-Romance. It is well known that Italian definite articles are governed by allomorphic rules. They propose that this is not strictly speaking allomorphy because all Italian articles share a common underlying form that varies in shape according to phonological conditions. Their major claim is that the articles underlying form and their morphological exponents have remained virtually unchanged since Medieval Italo-Romance and the only important changes have been to the phonological and morphosyntactic processes that affect their surface realisation.

In their Chapter they bring together diachrony, many modern dialects and Standard Italian under a shared, phonologically, and morphologically unified decompositional analysis.

In all the medieval and modern dialects investigated, the same phonological objects expone the same syntactic features, and the variation in surface forms are merely the effect of the micro-parametric differences in the phonological and morphosyntactic processes that interpret the same UR (underlying representation) according to its phonological and syntactic context. Their analysis includes dialectal forms with 'unexpected' gemination before vowelinitial stems (never previously analysed) and indefinite/mass articles in Southern Italo-Romance.

In the framework called a Word Syntax (i.e., morphological linearity is derived from hierarchical organization), they show that in these dialects' indefiniteness is marked by an indefinite bare marker 'de', in SpecDP within a layered DP structure (spelled out as P, such as 'de', only in the Medieval period). This bare marker acts as a licensor for a realized initial geminate of the masculine singular MASS and the indefinite feminine plural articles V-initial in the indefinite morphosyntactic environments. In C-initial position the φ-features of the indefinite articles geminate the onset of the following indefinite nouns via the Raddoppiamento Sintattico triggered by these two indefinite markers.

Gilbert Puech in his Chapter 5 ("Emergence of Maltese morpho-phonological profile") explores the Arabic introduced on the islands of Malta and Gozo, close to Sicily, in the High Middle Ages, and he endorses the view that Maltese is an offspring of Proto-Western Arabic, although Maltese has prosodic and syllabification-related features in common with pre-Hillalian Levantine dialects, and on the grounds that it shares with continental Maghrebi dialects morphological innovations which identify this group.

In his Chapter he focused on two aspects of Maltese morpho-phonology: syllabic structure and stress, and loss of 'back' consonants. The theoretical framework is structured by UG principles. He shows how emphatic consonants of Arabic Maltese were lost in the late Middle Ages and guttural consonants in modern times. This triggered substantial changes in Maltese phonology, and explains why Maltese, which is an Arabic dialect, now sounds very differently from the other dialect of the Arabic space. Once cut off from its Arabic and Islamic roots, new vocabulary was borrowed from European languages. The bilingualism with Italian, then with English impacted the morphophonological and morpho-lexical profile of modern Maltese.

Guillaume Enguehard analyzes in his Chapter 6 ("Sonority and Reduplication: an attempt to reduce the Sonority Condition to a Templatic Condition"), within Government Phonology theory, the reduplicative mechanisms in Gothic, Sanskrit, and Nakanai which implement the melodic complexity regarding the sonority condition. According to him the size of the template (i.e., the quantity of skeletal positions) conditions the sonority of the reduplicated segments. Complex less sonorous templates are associated to larger templates. In this way he provides in this chapter a theoretical approach that relates the principle of sonority and the template. This has implications on the theory of sonority and allows for questioning the sonority driven syllabification principle, since it is generally assumed that the distinction between the complex segments and cluster types is generated by the Sonority Sequencing Principle. The proposal of Guillaume Enguehard, based on diagnostics of syllable structure, is inverse to this traditional assumption: for him sonority derives from structure, not the other way around. The constraints on the reduplicated segments derive from the representations.

Timothée Premat, Sophie Chouvion and Axelle Verner (a singer, mezzo-soprano) in their Chapter 7 ("Repetitions, Rests Insertion and Schwa in 16 th century French Polyphony: An Emergent Sub-Grammar in Fresneau's Songs?") focus on the relationship between text and music in the 16 th century song, and on hierarchical constituent structure matching grammar in textsetting. Each song combines two objects, the text, the linguistic object, and the song, the musical object (tune). The authors describe how these two objects interact in Henry Fresneau's songs. The textsetting refers to the correspondence between text and tune, making use of formal similarities between language and music. One main similarity consists in the hierarchical constituent structure. Like syllables, lexical items can be grouped into phonological or syntactic constituents, the notes in a tune can be bounded in groups, also nested in hierarchical larger groups. Henry Fresneau's 16 th century polyphonic songs show a specific interaction in textsetting. The authors establish the matchings generated by this grammar and model the emergence of Fresneau's song sub-grammar. This chapter has benefited from the collaboration of a singer, Axelle Verner (mezzo-soprano) specialized in restoring a 16 th century French pronunciation, as well as on the analysis of textsetting in ancient music, and their transmission to singers and performers.

Maximilien Guérin examines in his Chapter 8 ("Locative, Presentative and Progressive Construction in Atlantic Languages") the verbal morphology of Atlantic Languages. He particularly focuses on a prototypical locative construction also used as a presentative and/or progressive construction and the marker associated to it. His hypothesis is that this construction, characteristic of this family, is inherited from Proto-Atlantic, and that its marker derives from a demonstrative determiner through a process of grammaticalization. His theoretical framework moves from grammatical constructions (usage-based Grammar), a functionalist approach to Grammars. His model of grammar diverges from the one adopted in the previous chapters since it is not based on the principles of universal grammar, rather in this model grammar is not independent of use and meaning, thus changes result from the creation of new grammars and the loss of the previous grammatical stages, processes called grammaticalizations from which the evolution of grammars would arise.

Jean Léo Léonard in his Chapter 9 ("Paradigm Function Morphology applied to the Southern Finnic Dialect Network") describes Finnic inflectional systems within a diasystemic analysis. He combines in his morpho-phonological analysis two different approaches, Paradigm Function Morphology (PFM), templatic phonology and particle phonology. He points out two main issues, relevant to Finno-Ugrian linguistics, general typology, and morphophonology: the relationship between Finnic inflectional taxonomies and typological inflection systems, and the challenge for theoretical linguistics to model morphological complexity and dialectal variation. He gives an account of Finnic inflectional taxonomies within PFM through a set of morphophonological rules and graphs of stem variation in order to turn 'complexity' into a 'simplexity' model. It is important to recall the core assumption of the Paradigm Function Morphology used in Chapter 9, in this framework Morphology is an autonomous component of the grammar. It means that in the internal architecture of Grammar, Morphology has an independent component with its own architecture, based on lexemes, paradigms, and realizations; thus, a language's inflectional morphology is the result of its paradigm function.

John te Velde and Nora Vosburg in Chapter 10 ("Plautdietsch: A remarkable story of Language and change") examine an endangered diasporic language of limited diffusion, Plautdietsch (also called Mennonite Low German), which has undergone changes dues to different linguistic factors. It is a religious minority originally from the Netherlands and Belgium, spoken as a primary language, and as heritage language. In the early formative years in the Vistula Delta, several phonological changes occurred that significantly changed the character of the original language, Dutch. It is composed by a cluster of dialects spoken among Mennonites in Europe, South America, and North America (USA, Canada, Mexico). Despite the actual distances, these varieties are still mutually intelligible. In this chapter they present interesting data from two Mennonite groups, one in Oklahoma, the other in Kansas. Despite the geographical distance for over a century their Plautdietsch varieties show a strikingly resemblance. Turning to formal aspects of syntax and phonology, they present evidence of the striking resilience of Plautdietsch dialect since earliest documentation. In particular, they found that the syntax of Plautdietsch varieties has remained intact as found in the other West Germanic languages with only very limited evidence of any shifts.

These ten contributions could be grouped in thematic sub-headings 'Syntax', 'Morphology', 'Morphosyntax', 'Phonology', 'Typology', but from some of them a common point of view emerges: morphological properties are dependent on syntax, and syntax, phonology and morphology are together responsible for interpretative properties providing classification criteria to typology. In some of these contributions also emerges the hypothesis that morphology derives basically from the application of phonology to syntactic structures. However, some contributions have stated that Morphology is an independent component. Thus, an important question is left open: what place should be respectively given to Syntax and Morphology in the grammatical derivations? For some researchers, some operations are part of the syntax, while others see them as independent and post-syntactic mechanisms, and for some researchers, phonology creates linearity independently of syntax, within morphemes.

The problem of the Syntax/Morphology and Phonology/Morphology interface is one of the major current issues in grammatical research. The themes proposed in this book particularly invite us to reflect.

Paris, the 14 th of Mai 2021, Michela Russo

Michela Russo. Introduction. Russo Michela (éd.). The Emergence of Grammars. A Closer Look at Dialects between Phonology and Morphosyntax. New York: Nova Sciences, Nova Sciences, pp.7-14., 2021, 978-1-53619-888-1. hal-03363237