General and specific effects of foundation species on the structure of alpine-fellfield communities in the Taisetsu Mountains, northern Japan

The great majority of studies on foundation species focused on a single dominant plant in a community, while more than one such species can often coexist and compete for space and limited resources. Morphologically different coexisting foundation species can create diverse niches occupied by different subsets of beneficiary species. To test this hypothesis, we sampled alpine plant communities at exposed fellfields in alpine zone, in the Taisetsu Mountains (Hokkaido, northern Japan), with coexisting putative foundation species Pinus pumila (evergreen shrub) and Diapensia lapponica (evergreen cushion-forming shrub), and analyzed their spatial relationships with other plants. Preliminary vegetation survey indicated that fruticose lichens and Loiseleuria procumbens (evergreen mat-forming shrub) might also act as a foundation species; thus, we included them in our analyses. The coexisting foundation species had both general as well as specific effects on plant community structure. Namely, almost all the members of the community aggregated spatially with lichens, while the other foundation species were spatially segregated from each other. These foundation plants associated with different members of the community, thus showing species-specific effects on the community structure. Blooming species showed even stronger patterns of species-specific spatial associations, suggesting that foundation species had beneficial effects on their associated species. We conclude that the focus on coexisting foundation species can reveal important details of community structure which would be hidden if we treated all species as equal members of the community. Studying the effects of coexisting foundation species could greatly advance our understanding of how species diversity functions in plant communities.