Environmental filtering, predominance of strong competitor trees and exclusion of moderate-weak competitor trees shape species richness and biomass.

Strong competitor (i.e. big-sized) trees are globally crucial for promoting aboveground biomass. Still, we do not fully understand the simultaneous influences of different levels of competitor (i.e. strong, moderate, medium and weak) trees at stand level in shaping forest diversity and biomass along a climatic gradient. We hypothesized that few strong competitor trees shape the positive relationship between tree species richness and aboveground biomass better than moderate, medium and weak competitor trees along a climatic gradient. Using the forest inventory data (i.e. tree diameter, height and crown diameter), we quantified strong (i.e. 99th percentile; top 1%), moderate (i.e. 75th percentile; top 25%), medium (i.e. 50th percentile) and weak (i.e. 25th percentile) competitor trees as well as species richness and aboveground biomass of 248 plots (moist temperate, semi-humid, and semi-arid forests) across 12 sites in Iran. The main results from three piecewise structural equation models (i.e. tree diameter, height and crown based models) showed that, after considering the simultaneous fixed effects of climate and random effects of sites or forest types variation, strong competitor trees possessed strong positive effects on tree species richness and biomass whereas moderate, medium and weak competitor trees possessed negligible positive to negative effects. Also, different levels of competitor trees promoted each other in a top-down way but the effects of strong competitor trees on moderate, medium and weak competitor trees were relatively weak. This study suggests that the simultaneous interactions of different tree sizes at stand level across forest sites should be included in the integrative ecological modeling for better understanding the role of different levels of competitor trees in shaping positive forest diversity - functioning relationship in a changing environment.

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Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.