Compounded perturbations in coastal areas: contrasting responses to nutrient enrichment and the regime of storm-related disturbance depend on life-history traits

1. Natural systems are exposed to compounded perturbations, whose changes in temporal variance can be as important as those in mean intensity for shaping the structure of assemblages. Specifically, climate-related physical disturbances and nutrient inputs due to natural and/or anthropogenic activities occur concomitantly, but experimental tests of the simultaneous effects of changes in the regime of more than one perturbation are generally lacking. Filling this gap is the key to understand ecological responses of natural assemblages to climate-related change in the intensity and temporal patterning of physical disturbance combined with other global stressors. 2. Responses to factorial manipulations of nutrient enrichment, mean intensity and temporal variability in storm-like mechanical disturbance were examined, using benthic assemblages of tide-pools as model system. 3. Response variables were mean abundance values and temporal variances of taxa with different life-traits. Consistent negative effects of disturbance intensity were observed for the mean cover of long-living taxa (algal canopies and the polychaete Sabellaria alveolata), whose temporal fluctuations were also reduced by more severe mechanical stress. More resilient taxa (ephemeral algae, mostly green of the genus Ulva) increased under enriched conditions, particularly when low-intensity events were irregularly applied over time. Opposite effects of disturbance intensity depending on nutrient availability occurred on filamentous algae (e.g. red of the genus Ceramium). This was probably due to the fact that, although nutrient enrichment stimulated the abundance of both algal groups, when this condition was combined with relatively mild physical disturbance the competitively superior ephemeral green algae tended to become dominant over filamentous red algae. The same did not occur under high intensity of disturbance since it likely damaged large, foliose fronds of Ulva-like forms more than small, filamentous fronds of Ceramium-like forms. Grazers were positively affected by nutrients, likely responding indirectly to more food available. 4. A direct relationship between the mean abundance of most organisms and their temporal fluctuations was documented. However, all organisms persisted throughout the study, even under experimental conditions associated to the largest temporal variation in their abundance, likely due to their ability to resist to/quickly recover from, the applied perturbations. Therefore, in systems with great recovery abilities of dominant organisms (e.g. rocky intertidal, grasslands), effects of traits of the regime of disturbance and nutrient enrichment may modulate the fluctuations of populations not through the elimination and substitution of species, but through changes in relative abundances of the same species. This contrasts with the theory that temporal variation in abundance would be directly related to the risk of local extinction. Present findings enable more accurate predictions of the consequences of climatic and non-climatic scenarios on the biodiversity of marine and terrestrial systems sharing analogous functional traits of organisms. Future more intense physical disturbances are expected to exert negative effects on slow-growing/recovering species (e.g. habitat-formers) irrespectively of the temporal patterning of the same disturbances and nutrient inputs. On the contrary, more resistant species (e.g. encrusting algae on rocky shores or below-ground vegetation in grasslands) are expected to benefit from intense physical disturbance. Species whose abundance is more directly related to the availability of nutrients (e.g. filamentous and ephemeral algae or herbs) are expected to generally increase under enriched conditions, but their ability to eventually become dominant would depend on their ability to grow fast and attain cover large enough to overwhelm any possible control of concomitant disturbance intensity on their abundance. If, such as in the present examined system, virtually all organisms can persist, over the temporal scale of the experiment, under any combination of physical disturbance and nutrient availability, the resulting overall diversity is not predicted to change drastically. Nevertheless, low-intensity events evenly distributed and high-intensity events irregularly distributed appear as the conditions supporting the highest richness of taxa, independently of the availability of nutrients.POFC-COMPETEQuadro de Referencia Estrategico Nacional (QREN)FCTErasmus grant of the European MSc in Marine Biodiversity and Conservation (EMBC)[PTDC/MAR/111223/2009][PEst-C/MAR/LA0015/2011][PEst-C/MAR/LA0015/2013][BGCT/MAR/LA0015-2014