In relation to plant species (Fig. 4), average and total herb and

tree species richness were both highest in pine plantations (in total 31 and 11 species, respectively), followed by mixed forests (in total 26 and 10 species, respectively), while average and total shrub diversity was highest in birch (in total 6 species) forests. The lowest vascular plant species richness in all three layers both on average and in total was recorded in oak forests (in total 19 herb, 2 shrub and mTOR inhibitor 4 subdominant tree species). Analysis of the community composition of carabid assemblages (Fig. 5) reveals that the pine plantation and oak forest harbour distinct communities relative to the other forest types. Furthermore, it is apparent that oak, pine and mixed forests show greater heterogeneity in community composition and therefore a higher species turnover between plots than the other forest types. By contrast, birch and larch forest plots show relatively little variation in the species composition. The environmental parameters investigated in this study (Table 1) exerted only a limited amount of control over the beetle distribution patterns, with the first two RDA axes explaining only 16.2% and 5.9% of species variation, respectively. Both canopy cover and dry weight of the litter layer exerted

some influence, with larch and birch forests being characterised by a high amount of litter and open canopies (Fig. 6). Oak and Ureohydrolase pine forests were both characterised by closed canopies, but oak forest litter buy Ruxolitinib had a lower relative dry weight. Mixed forests were most heterogeneous in relation to environmental parameters, mirroring the high levels of heterogeneity observed in carabid species composition between samples in this forest type. Most carabid species are clustered towards the centre of the RDA plot. The abundances of some of these species are likely too low to result in a clear environmental response pattern, while

other species may be unaffected by the recorded variables or prefer intermediate environmental settings. However, all five dominant species are clearly associated with distinct habitat conditions. C.vladimirskyi associates strongly with high canopy cover and low leaf litter mass that characterises oak forest samples, while C. crassesculptus also associates strongly with high canopy cover, but only intermediate leaf litter mass and low ground cover. By contrast, P.acutidens has a strong association with open canopies and a high leaf litter mass. P.adstrictus and C.manifestus associate with intermediate values of these parameters. Furthermore, Synuchus sp. and Harpalus coreanus (Tschitscherin, 1895) are notable due to their association with higher ground vegetation cover values.