Vertical zonation is the main distribution pattern of littoral assemblages on rocky shores at a regional scale

Vertical variation in the distribution of rocky shore assemblages is greater than horizontal variation, as shown by univariate and multivariate analysis performed with data obtained along 1000 km of shoreline and covering from the upper supralittoral to the upper infralittoral zone (−1 m). Consequently, vertical littoral zonation is a consistent pattern at a regional scale within the same biogeographical zone. While their distribution varies at the same shore height, marine species and assemblages from rocky shores show a specific vertical sequence known as zonation. A key question in ecology is how consistent is zonation along large spatial scales. The aim of this study is to show distribution patterns of littoral assemblages at a regional scale and to identify the most relevant abiotic factors associated to such patterns. The study is based on a detailed and extensive survey at a regional scale on a tideless rocky shore. Benthic macroflora and macrofauna of 750 relevés were described along the vertical axis of 143 transects distributed across the shoreline of Catalonia (NW Mediterranean). The Detrended Correspondence Analysis (DCA) first axis is highly related to the height on the shore: species, relevés, and assemblages grade from lower to upper height (infralittoral to supralittoral). As observed in nature, different assemblages co-occur at the same height at different sites, which is shown along DCA second axis. The abiotic variables that best explain the assemblage distribution patterns are: height (75% of the model inertia), longitude (14.6%), latitude (7.2%) and transect slope (2.9%). The Canonical Correspondence Analysis (CCA) first axis is related to height on the shore and explains four times more variance than CCA second axis, which is related to the horizontal gradient. Generalized Lineal Model (GLM) results show that height on the shore is the factor explaining most of the variance in species presence. Most studied species show distribution patterns related to latitude and longitude, but always in a much smaller proportion than to height.