In rocky intertidal habitats, the pronounced increase in environmental stress from low to high elevations greatly affects community structure, that is, the combined measure of species identity and their relative abundance. Recent studies have shown that ecological variation also occurs along the coastline at a variety of spatial scales. Little is known, however, on how vertical variation compares with horizontal variation measured at increasing spatial scales (in terms of sampling interval). Because broad-scale processes can generate geographical patterns in community structure, we tested the hypothesis that vertical ecological variation is higher than fine-scale horizontal variation but lower than broad-scale horizontal variation. To test this prediction, we compared the variation in community structure across intertidal elevations on rocky shores of Helgoland Island with independent estimates of horizontal variation measured at the scale of patches (quadrats separated by 10s of cm), sites (quadrats separated by a few m), and shores (quadrats separated by 100s to 1000s of m). The multivariate analyses done on community structure supported our prediction. Specifically, vertical variation was significantly higher than patch- and site-scale horizontal variation but lower than shore-scale horizontal variation. Similar patterns were found for the variation in abundance of foundation taxa such as Fucus spp. and Mastocarpus stellatus, suggesting that the effects of these canopy-forming algae, known to function as ecosystem engineers, may explain part of the observed variability in community structure. Our findings suggest that broad-scale processes affecting species performance increase ecological variability relative to the pervasive fine-scale patchiness already described for marine coasts and the well known variation caused by vertical stress gradients. Our results also indicate that experimental research aiming to understand community structure on marine shores should benefit from applying a multi-scale approach.

Environmental stress affects species richness and diversity in communities, but the precise form of the relationship is unclear. We tested an environmental stress model (ESM) that predicts a unimodal pattern for total richness and diversity in local communities across the full stress gradient where a regional biota can occur. In 2008, we measured richness and diversity (considering all macrobenthic species) across the entire intertidal range on three rocky shores on Helgoland Island, Germany. Intertidal elevation is known to be positively related to abiotic stress. Since Helgoland is between the northern and southern biogeographic boundaries for the cold-temperate NE Atlantic intertidal biota, it exhibits low stress levels for this biota at low elevations and high stress at high elevations because of long ([6 h) emersion times. Thus, we predicted a unimodal trend for richness and diversity across elevation. On all three shores, richness increased from high to middle elevations, but remained similar between middle and low elevations. Diversity followed the same trend on one shore and different trends (although also nonunimodal) on the other two. Evenness explained the trend differences between richness and diversity. Overall, our study yielded little support for the ESM. Reasons for richness and diversity not decreasing at low elevations may be related to influences of mostly subtidal species, Helgoland's intertidal range, or sampling resolution. Our study also suggests that the ESM must be developed further to differentiate between richness and diversity. We offer recommendations to improve future ESM research using intertidal systems.