Disentangling direct and indirect effects of island area on plant functional trait distributions

Aim Species diversity on islands generally increases with island area. This might arise either from direct effects of island area via neutral assembly processes or from indirect effects via habitat and structural differences between islands that scale positively with island area. Here, we tested whether community-weighted functional trait means of woody plants are directly or indirectly affected by island area to elucidate how functional traits mediate the assembly on differently sized islands. Location Twenty-eight tropical islands (25 m(2) - 12,000 m(2)) in the Raja Ampat archipelago, Indonesia. Taxon Woody angiosperms. Methods Studied islands had a shared geological history but differed in terms of area, habitat quality expressed by soil depth, forest structure expressed by tree basal area and degree of isolation. Traits studied were seed and fruit mass, tree height, wood density, leaf mass per area, leaf nitrogen concentration and chlorophyll content (estimated from chlorophyll-meter units) and summarised as community-weighted means (CWM) for each island. Using liner regression, we tested whether CWMs were correlated to island area and basal area and structural equation models (SEMs) to test on direct and indirect effects of island area, basal area, soil depth and isolation on trait distributions. Results CWM of seed mass, tree height and chlorophyll content increased with both island area and basal area, whereas leaf nitrogen concentration decreased with increasing basal area. Fruit mass was not correlated to island area and basal area. SEMs revealed that the shifts in tree height, wood density, leaf nitrogen concentration and chlorophyll content were caused directly by basal area, which in turn was directly and positively affected by both island area and soil depth. Differences in seed mass among islands were explained by combined effects of basal area, island area and isolation, whereas fruit mass was only explained by isolation. Main conclusions Trait values shifted systematically across islands of different sizes. Being small and having light seeds are prevailing trait combinations for establishing on small islands with simple forest structure. For establishment on larger islands with more complex forest structures, species are taller, have heavier seeds, higher chlorophyll content and lower leaf N concentrations. We conclude that mechanisms affecting CWM on islands directly link to ecological differences between islands like forest structure - and only indirectly to island area.

Automated summary

Species diversity on islands generally increases with island area. Traits of woody plants, such as tree height, wood density, leaf nitrogen concentration and chlorophyll content, are mainly affected by basal area, which in turn is influenced by both island area and soil depth. Seed and fruit mass variations among islands are explained by a combination of basal area, island area, and isolation.