Size matters sometimes: wall height and the structure of subtidal benthic invertebrate assemblages in south-eastern Australia and Mediterranean Spain

Aim Variation in the structure of shallow subtidal invertebrate assemblages was examined over three spatial scales; within reef, between reef and between continents. We sought to provide a context from which to examine and interpret ecological processes between continents. In addition, we predicted that variation in pattern would increase as the scale of examination increased. Location Reefs near Wollongong and within Jervis Bay in south-eastern Australia and Mediterranean reefs on the Costa Brava (Catalonia), north-eastern Spain. Methods We compared assemblages on vertical rock walls of two heights - short (< 2 m) and tall (> 3 m) in two temperate regions over the same depth range. Specifically we examined the diversity and cover of invertebrates, the cover and biomass of foliose and crustose algae, the size of invertebrate colonies and the biomass of urchins on short and tall walls (n = 3) at each of two locations in each country. Results Foliose algae dominated rock walls in Spain and although invertebrate cover was high, colonies were generally very small. Two urchin species were commonly encountered on rock walls in Spain, Arbacia lixula and Paracentrotus lividus; their biomass was relatively low and did not differ significantly between short and tall walls. These findings contrasted strongly with south-eastern Australia, where foliose algae were almost completely absent. A single urchin species, Centrostephanus rodgersii occurred with extremely high biomass on short walls, which were dominated by grazer-resistant crustose calcareous algae. In contrast, the biomass of this urchin was low on tall walls, which were dominated by invertebrates, usually exceeding 95% in cover. Invertebrate colonies were significantly larger on both short and tall walls in south-eastern Australia relative to the Mediterranean. Findings within a country were consistent between the replicate rock walls and between locations. In contrast to our prediction, however, there was significant variation among walls within a location, but not among locations within a continent. Temporal variation in the structure of these assemblages was not examined, but appears limited. Main conclusions We conclude that submarine topography, i.e. the presence of short or tall rock walls, as a function of rock type and structure, has a marked impact on community structure in south-eastern Australia, but made little difference to the structure of the assemblage in Mediterranean Spain. The differences in structure we observed between walls of different heights in Australia were correlated with differences in the biomass of urchins and they appear to be major determinants of assemblage structure. Interactions among species are often reported from disparate parts of the globe with little or no reference to the structure of the assemblage of which they are a part; we contend that this will hinder interpretation. Our data are consistent with the organisms in these two regions experiencing distinct selection pressures; for example high levels of urchin grazing activity in south-eastern Australia, and shading and whiplash associated with an algal canopy in the Mediterranean. It may not be appropriate to contrast processes operating at very large (intercontinental) scales unless context can be established with a clear understanding of ecological pattern.