Marine macrophytes directly enhance abundances of sandy beach fauna through provision of food and habitat

Beach-cast wrack is a prominent feature of beaches of south-western Australia. We examined the fauna of these beaches to explore the generalisation [Polis, G.A., Hurd, S.D., 1995. Extraordinarily high spider densities on islands: flow of energy from the marine to terrestrial food webs and the absence of predation. Ecology 92, 4382e4386] that beach-cast wrack from highly productive marine ecosystems subsidises low productivity of terrestrial ecosystems, to establish whether this generalisation is relevant to oligotrophic marine systems. We sampled three beaches with high and three beaches with low volumes of beach-cast wrack to determine if: (1) the presence of wrack influences the abundance of macroinvertebrates; (2) wrack acts as a food source for beach macroinvertebrates; and (3) the influence of wrack varies between zones above the high water mark. We measured wrack volume and composition, sediment characteristics, the abundance of different epibenthic and infaunal macroinvertebrates taxa, and d13C and d15N of macrophytes and macroinvertebrates. The mean volume of wrack on high-wrack beaches was 0.27e1.07 m3 wrack m 2 compared to 0.00e0.09 m3 wrack m 2 on low-wrack beaches. There were no significant differences in sediment grain size, moisture content or loss on ignition between the two types of beaches or zones. Epibenthic fauna and infauna were consistently abundant on high-wrack beaches (20e291 and 0.5e3.5 individuals 0.64 m 2, respectively), but either absent or extremely rare in low-wrack beaches (0e3 and 0e0.1 individuals 0.64 m 2, respectively). Within high-wrack beaches, there were no significant differences in the abundance of epifauna or infauna among beaches or between zones. The d13C values of macroinvertebrates at all sites were most similar to red and brown algae, with the exception of beetles from two beaches, which were closest to seagrasses. Mixing model (Isosource) results for mesograzing amphipods and dipteran flies suggested carbon was assimilated mostly from the seagrass Posidonia spp., the dune grass Spinifex longifolia and red algae for amphipods and from brown algae, red algae and dune vegetation for dipteran flies. We conclude that the presence of marine-derived wrack plays a major role in subsidising production of macroinvertebrates on beaches of south-western Australia. We suggest that marine subsidies can play a role in supporting terrestrial production, even in oligotrophic marine environments.