Bioengineer effects on understory species richness, diversity, and composition change along an environmental stress gradient: Experimental and mensurative evidence

Canopy-forming bioengineer species are commonly assumed to increase local species richness and diversity. We tested this notion by investigating the effects of fucoid seaweed canopies on understory communities along rocky intertidal elevation gradients in Atlantic Canada. Such gradients exhibit increasing thermal extremes and variation from low to high elevations, and are broadly used in stress gradient studies. A manipulative experiment created canopy and no-canopy treatments at the low, middle, and high intertidal zones, eliminating all species (except fucoid canopies) from replicate quadrats. After recolonization, overall richness and diversity (considering all primary producers and consumers) were higher under canopies than uncovered by canopies at the high and middle zones, but no effects occurred at the low zone. Similarly, species composition was affected by canopies at the high and middle zones, but not at the low zone. A mensurative study that surveyed the full range of canopy cover (0–100%) using nearly five times more quadrats from pristine areas yielded the same results: richness and diversity increased with canopy cover at the high and middle zones (approaching stabilization toward high cover values), but no effects occurred at the low zone. Lack of canopy effects at low elevations is related to mild habitat conditions, which canopies are unable to modify, while positive effects at higher elevations relate to the capacity of canopies to ameliorate harsh conditions. This is the first time that a combined experimental and mensurative approach shows that the same bioengineer species affect overall species richness, diversity, and composition differently along a stress gradient. Overall, protecting canopy-forming bioengineers to preserve local biodiversity should be most effective in stressful environments.