Heterogeneity–diversity relationships differ between and within trophic levels in temperate forests

The habitat heterogeneity hypothesis predicts that biodiversity increases with increasing habitat heterogeneity due to greater niche dimensionality. However, recent studies have reported that richness can decrease with high heterogeneity due to stochastic extinctions, creating trade-offs between area and heterogeneity. This suggests that greater complexity in heterogeneity–diversity relationships (HDRs) may exist, with potential for group-specific responses to different facets of heterogeneity that may only be partitioned out by a simultaneous test of HDRs of several species groups and several facets of heterogeneity. Here, we systematically decompose habitat heterogeneity into six major facets on ~500 temperate forest plots across Germany and quantify biodiversity of 12 different species groups, including bats, birds, arthropods, fungi, lichens and plants, representing 2,600 species. Heterogeneity in horizontal and vertical forest structure underpinned most HDRs, followed by plant diversity, deadwood and topographic heterogeneity, but the relative importance varied even within the same trophic level. Among substantial HDRs, 53% increased monotonically, consistent with the classical habitat heterogeneity hypothesis but 21% were hump-shaped, 25% had a monotonically decreasing slope and 1% showed no clear pattern. Overall, we found no evidence of a single generalizable mechanism determining HDR patterns.