We also examined relationships between benthos and sediment at scales of 1–2 m (among cores/quadrats within sites), 10 s of metres (among sites) and kilometres (among bays) within each habitat. Because the physical complexity of the substratum is increased by the presence of pneumatophores, leaf litter and macro-algae and these features were patchy in the different forests, it was expected that two habitats (MH1 and MH2) would show greater variability among sites and bays than would bare mud (MH3). The benthos varied at the larger scales, with significant variability among bays, in all habitats. The sediment did not, however, differ significantly among bays in MH3-the permanently shaded habitat, probably due to extremely large site-to-site variability.