Species–area relationships (SARs) characterize the spatial distribution of species diversity in community ecology, but the biological mechanisms underlying the SARs have not been fully explored. Here, we examined the roles of dispersal limitation and habitat heterogeneity in shaping SARs in two large‐scale forest plots. One is a 24‐ha subtropical forest in Gutianshan National Nature Reserve, China. The other is a 50‐ha tropical rain forest in Barro Colorado Island, Panama. Spatial point pattern models were applied to investigate the contributions of dispersal and habitat heterogeneity and their interactions to the formation of the SARs in the two sites. The results showed that, although dispersal and habitat heterogeneity each could significantly contribute to the SARs, each alone was insufficient to explain the SARs. Their joint effects sufficiently explained the real SARs, suggesting that heterogeneous habitat and dispersal limitation are two predominant mechanisms for maintaining the spatial distributions of the species in these two forests. These results add to our understanding of the ecological processes underlying the spatial variation of SARs in natural forest communities.

Large old trees are keystone ecological structures that provide vital ecosystem services to humans. However, there are few large-scale empirical studies on patterns of diversity and density of large old trees in human-dominated landscapes. We present the results of the first nationwide study in China to investigate the patterns of diversity and density of large old trees in human-dominated landscapes. We collated data on 682,730 large trees ≥100 years old from 198 Chinese regions to quantify tree species diversity, tree density and maximum tree age patterns. We modelled the effects of natural environmental variables (e.g. climate and topography) and anthropogenic variables (e.g. human population density and city age) on these measures. We found a low density of large old trees across study regions (0.36 trees/km2), and large variation in species richness among regions (ranging from 1 to 232 species). More than 95% of trees were <500 years old. The best fit models showed that: (1) Species diversity (species richness adjusted by region size) was positively associated with mean annual rainfall and city age; (2) Density of clustered trees, which are mostly remnants of ancient woods, was negatively influenced by human population density and rural population (% of total population). In contrast, the density of scattered trees, which are mostly managed by local people, was positively correlated with mean annual rainfall and human population density. To better protect large old trees in cities and other highly-populated areas, conservation policy should protect ancient wood remnants, mitigate the effects environmental change (e.g. habitat fragmentation), minimize the negative effects of human activities (e.g. logging), and mobilize citizens to participate in conservation activities (e.g. watering trees during droughts).

Spatially explicit consideration of species distribution can significantly add to our understanding of species coexistence. In this paper, we evaluated the relative importance of habitat heterogeneity and other clustering processes (e.g., dispersal limitation, collectively called the non‐habitat clustering process) in explaining the spatial distribution patterns of 341 tree species in three stem‐mapped 25–50 ha plots of tropical, subtropical, and temperate forests. Their relative importance was estimated by a method that can take one mechanism into account when estimating the effects of the other mechanism and vice versa. Our results demonstrated that habitat heterogeneity was less important in explaining the observed species patterns than other clustering processes in plots with flat topography but was more important in one of the three plots that had a complex topography. Meanwhile, both types of clustering mechanisms (habitat or non‐habitat) were pervasive among species at the 50‐ha scale across the studied plots. Our analyses also revealed considerable variation among species in the relative importance of the two types of mechanism within each plot and showed that this species‐level variation can be partially explained by differences in dispersal mode and growth form of species in a highly heterogeneous environment. Our findings provide new perspectives on the formation of species clustering. One important finding is that a significant species–habitat association does not necessarily mean that the habitat heterogeneity has a decisive influence on species distribution. The second insight is that the large species‐level variation in the relative importance of the two types of clustering mechanisms should not be ignored. Non‐habitat clustering processes can play an important role on species distribution.