Carabids benefit more from pine stands with added understory or second story of broad-leaved trees favored by climate change than from one-storied pine stands

The released excess anthropogenic nitrogen and carbon produces habitat enrichment, as exemplified by the modification of one-storied pine stands by introducing deciduous species into the understory or second story. In this study, we discuss the validity of pine stand modification by pitfall-trapping epigeic carabid beetles. Two hypotheses were formulated: (1) proportion of late-successional species is higher in assemblages inhabiting pine stands with understory or a second story than in one-storied pine stands; (2) plant litter composition affects carabid beetle assemblages more than other environmental variables. Additionally, characteristic carabid species of the respective pine stand types were identified. GLMM analysis revealed a higher proportion of late-successional species in pine stands with understory or a second story than in one-storied ones. NMDS separated those pine stand types. RDA analysis indicated that pine litter and humus had the strongest effect on carabid beetle assemblage structure in one-storied stands, being drier and thicker in this stand type than in the others. Indicator value analysis identified two characteristic non-forest species in one-storied stands The study revealed that the introduction of understory and particularly a second story into pine stands increased carabid beetle diversity and the proportion of late-successional species, confirming the validity of pine stand modification.