Canopy tree species determine herb layer biomass and species composition on a reclaimed mine spoil heap

According facilitative models of succession, trees are great forest ecosystem engineers. The strength of tree stand influences on habitat were tested in rather homogenous conditions where heterogeneity of site condition was not an important influence. We hypothesized that canopy composition affects total aboveground vascular herb layer biomass (THB) and species composition of herb layer plant biomass (SCHB) more significantly than primary soil fertility or slope exposure. The study was conducted in 227 randomly selected research plots in seven types of forest stands: pure with Alnus glutinosa, Betula pendula, Pinus sylvestris, Quercus petraea and Robinia pseudoacacia, and mixed with dominance of Acer pseudoplatanus or Betula pendula located on hilltop and northern, eastern, western, and southern slopes on a reclaimed, afforested post-mining spoil heap of the Bełchatów Brown Coal Mine (Poland). Generalized linear models (GLZ) showed that tree stand species were the best predictors of THB. Non-parametric variance tests showed significantly higher (nearly four times) THB under canopies of A. glutinosa, R. pseudoacacia, B. pendula and Q. petraea, compared to the lowest THB found under canopies of P. sylvestris and mixed with A. pseudoplatanus. Redundancy Analysis (RDA) showed that SCHB was significantly differentiated along gradients of light-nutrient herb layer species requirements. RDA and non-parametric variance tests showed that SCHB under canopies of A. glutinosa, R. pseudoacacia and mixed with A. pseudoplatanus had large shares of nitrophilous ruderal species (32%, 31% and 11%, respectively), whereas SCHB under B. pendula, Q. petraea, mixed with B. pendula and P. sylvestris were dominated by light-demanding meadow (49%, 51%, 51% and 36%, respectively) and Poaceae species. The results indicated the dominant role of tree stand composition in habitat-forming processes, and although primary site properties had minor importance, they were also modified by tree stand species.