Effects of habitat area and spatial configuration on biodiversity in an experimental intertidal community

Isolating the effects of fragmentation per se (i.e., spatial configuration of habitat patches) on species richness is an ongoing challenge as habitat configuration often covaries with the amount of habitat. Consequently, there is a lack of experimental evidence for configurational effects on species richness in the whole landscape. Here, we developed a novel experimental system for testing the independent and interactive effects of habitat area and configuration on tropical intertidal species richness. Our results confirmed the expectation that average species richness would increase monotonically with habitat area. More intriguingly, we found mixed evidence for a non-monotonic relationship between species richness and fragmentation per se, with the highest richness at intermediate fragmentation configuration, that is, when habitat tiles were placed in a "several-small" configuration. The effect of habitat configuration was not due to passive sampling (since area was controlled for), variation in total individual abundance, or niche specialization of species to different landscape configurations. We postulate that a combination of processes, including local negative density dependence and dispersal limitation, could give rise to the observed pattern. We emphasize the importance of considering configurational effects on biodiversity at broader spatial scales and for more experimental research to delve into the mechanisms driving the patterns seen here.