Indirect effects shape macroalgal epifaunal communities.

We tested the response of algal epifauna to the direct effects of predation and the indirect consequences of habitat change due to grazing and nutrient supply through upwelling using an abundant intertidal rhodophyte, Gelidium pristoides. We ran a mid-shore field experiment at four sites (two upwelling sites interspersed with two non-upwelling sites) along 450 km of the south coast of South Africa. The experiment was started in June 2014 and ran until June 2015. Four treatments (predator exclusion, grazer exclusion, control, and procedural control) set out in a block design (n = 5) were monitored monthly for algal cover for the first 6 months and every 2 months for the last 6 months. Epifaunal abundance, species composition, algal cover, and algal architectural complexity (measured using fractal geometry) were assessed after 12 months. Predation had no significant effect on epifaunal abundances, while upwelling interacted with treatment. Grazing reduced the architectural complexity of algae, with increased fractal dimensions in the absence of grazers, and also reduced algal cover at all sites, though the latter effect was only significant for upwelling sites. Epifaunal community composition was not significantly affected by the presence of herbivores or predators but differed among sites independently of upwelling; sites were more similar to nearby sites than those farther away. In contrast, total epifaunal abundance was significantly affected by grazing, when normalized to algal cover. Grazing reduced the cover of algae; thus, epifaunal abundances were not affected by the direct top-down effects of predation but did respond to the indirect effects of grazing on habitat availability and quality. Our results indicate that epifaunal communities can be strongly influenced by the indirect consequences of biotic interactions.

Cheating the Locals: Invasive Mussels Steal and Benefit from the Cooling Effect of Indigenous Mussels.

The indigenous South African mussel Perna perna gapes during periods of aerial exposure to maintain aerobic respiration. This behaviour has no effect on the body temperatures of isolated individuals, but when surrounded by conspecifics, beneficial cooling effects of gaping emerge. It is uncertain, however, whether the presence of the invasive mussel Mytilus galloprovincialis limits the ability of P. perna for collective thermoregulation. We investigated whether varying densities of P. perna and M. galloprovincialis influences the thermal properties of both natural and artificial mussel beds during periods of emersion. Using infrared thermography, body temperatures of P. perna within mixed artificial beds were shown to increase faster and reach higher temperatures than individuals in conspecific beds, indicating that the presence of M. galloprovincialis limits the group cooling effects of gaping. In contrast, body temperatures of M. galloprovincialis within mixed artificial mussel beds increased slower and exhibited lower temperatures than for individuals in beds comprised entirely of M. galloprovincialis. Interestingly, differences in bed temperatures and heating rates were largely dependent on the size of mussels, with beds comprised of larger individuals experiencing less thermal stress irrespective of species composition. The small-scale patterns of thermal stress detected within manipulated beds were not observed within naturally occurring mixed mussel beds. We propose that small-scale differences in topography, size-structure, mussel bed size and the presence of organisms encrusting the mussel shells mask the effects of gaping behaviour within natural mussel beds. Nevertheless, the results from our manipulative experiment indicate that the invasive species M. galloprovincialis steals thermal properties as well as resources from the indigenous mussel P. perna. This may have significant implications for predicting how the co-existence of these two species may change as global temperatures continue to rise.

Not all space is created equal: distribution of free space and its influence on heat-stress and the limpet Patelloida latistrigata.

For most marine benthic communities unoccupied primary substrata, or free space, is considered the principle limiting resource. Substratum temperatures, desiccation rates and hydrodynamic characteristics of free space, however, may vary depending on patch size and isolation and therefore potentially influence biotic processes. This paper investigates the relationship between small-scale changes in the availability and configuration of free space, heat stress and abundance of the small rocky intertidal gastropod Patelloida latistrigata within southeastern Australia. Using infrared thermography I show that heat stress of rocky intertidal communities increased linearly with increasing amounts of free space on three neighbouring shores during four separate sampling intervals from October 2009 to January 2010. Abundances of P. latistrigata generally declined with increasing availability of free space and the associated increases in heat stress. An experimental manipulation that altered the configuration but not the availability of free space demonstrated that both heat stress and P. latistrigata abundance are not affected by small-scale changes in the configuration of free space. The small-scale distribution of P. latistrigata, however, was significantly influenced by differences in the configuration of free space with limpets displaying bimodal distributions within areas characterised by unevenly distributed free space. Since the distribution of Patelloida varies depending on the configuration of free space but thermal properties at the scale of individual limpets do not then we might expect Patelloida to be responding to changes in other abiotic factors, such as hydrodynamic forces and desiccation rates, which may change with the configuration of free space. This study highlights the dynamic and usually unexamined relationship between abiotic stress and the availability and acquisition of resources by marine benthic invertebrates.