The seaweed Caulerpa racemosa on Mediterranean rocky reefs: from passenger to driver of ecological change.

Disentangling the ecological effects of biological invasions from those of other human disturbances is crucial to understanding the mechanisms underlying ongoing biotic homogenization. We evaluated whether the exotic seaweed, Caulerpa racemosa, is the primary cause of degradation (i.e., responsible for the loss of canopy-formers and dominance by algal turfs) on Mediterranean rocky reefs, by experimentally removing the invader alone or the entire invaded assemblage. In addition, we assessed the effects of enhanced sedimentation on the survival and recovery of canopy-forming macroalgae at a relatively pristine location and how their loss affects the ability of C. racemosa to conquer space. C. racemosa did not invade dense canopy stands or influence their recovery in cleared plots. Competition with C. racemosa could not explain the rarity of canopy-forming species at degraded sites. Removing the assemblages invaded by C. racemosa and preventing reinvasion did not trigger the transition from algal turfs to canopies, but it enhanced the cover of morphologically complex erect macroalgae under some circumstances. Once established, C. racemosa, enhancing sediment accumulation, favors algal turfs over erect algal forms and enables them to monopolize space. Our results show that introduced species that rely on disturbance to establish can subsequently become the main drivers of ecological change.

The spread of Caulerpa racemosa var. cylindracea in the Mediterranean Sea: an example of how biological invasions can influence beta diversity.

The present study aims to evaluate if invasion of Caulerpa racemosa var. cylindracea can affect both alpha and beta diversity in Mediterranean coastal system. The following hypotheses were tested: (i) measures of alpha diversity differ between invaded and non-invaded assemblages (ii) this pattern is consistent between different habitats, (iii) assemblages from habitats which are naturally very different tend to be more similar when invaded compared to non-invaded situations. Results showed that Mediterranean benthic assemblages invaded by C. racemosa var. cylindracea were characterized by lower values of alpha diversity than non-invaded assemblages; moreover, differences between deep (25 m) and shallow water (5m) assemblages appeared lower in invaded than in non-invaded areas, with a decrease of beta diversity. This homogenisation of habitats was related to a lack of several characteristic species and to a higher abundance of few opportunistic species in invaded areas, particularly turf-forming algae. Moreover, variability in taxa composition was lower in invaded than in non-invaded assemblages in both habitats.

Sediment disturbance and loss of beta diversity on subtidal rocky reefs.

How changes in environmental complexity and heterogeneity affect beta diversity is poorly known. We investigated patterns of beta diversity in subtidal assemblages of algae and invertebrates in the northwest Mediterranean in relation to inclination of the substratum and sedimentation. Vertical and horizontal substrata supported distinct assemblages under low, but not under heavy, ambient loads of sediment. To test the hypothesis that sediment reduced the dissimilarity between assemblages, sedimentation was increased experimentally in plots established on vertical and horizontal surfaces at sites experiencing low ambient levels of sedimentation. Patterns were compared to those occurring at unmanipulated sites and at sites exposed to heavy loads of sediment about 2 km apart. After one year, assemblages on vertical substrata were indistinguishable from those occurring on flat surfaces at manipulated sites and both converged toward those occurring at sites exposed to heavy loads of sediment. Control sites still supported distinct assemblages on vertical and horizontal substrata by the end of the experiment. Similar effects of sediment were observed on recovering assemblages in experimental clearings. These results show that sediment increased similarity in assemblages overriding the influence of habitat complexity on beta diversity at small and large spatial scales.

Variability of Mediterranean coralligenous assemblages subject to local variation in sediment deposition.

This study tested whether the development of coralligenous assemblages on horizontal and vertical surfaces differed between localities of high and low sediment deposition. The development and structure of these assemblages varied in predictable ways according to the level of sediment deposition. These differences were largely independent of the orientation of substratum. Turfs were more extensive in areas of high sediment deposition while erect and encrusting algae were most extensive in areas of low sediment deposition. Encrusting invertebrates characterised vertical surfaces and were most extensive in areas of high sediment deposition. These results are consistent with studies from other temperate regions of the globe, suggesting that effects of sedimentation on temperate coasts are substantial and widespread.

Threats to macroalgal coralligenous assemblages in the Mediterranean Sea.

Coralligenous habitat is one of the most important coastal systems of the Mediterranean Sea. This paper evaluates the main threats to macroalgal coralligenous habitats through a review of the relevant literature. Sedimentation, nutrient enrichment and biological invasions can cause severe alterations in the structure of coralligenous assemblages due to the regression of perennial structuring species and increases in populations of ephemeral algae. Under pristine conditions, mechanical disturbance seems to be easily mitigated by the recovery capability of coralligenous assemblages; however, such disturbances seriously affect coralligenous structure if they occur in concert with other stressors. Important synergetic effects among all the studied anthropogenic disturbances are also highlighted. The main consequences of the considered stressors are the loss of the complexity of macroalgal assemblages and the deterioration of both alpha and beta diversity.

Effects of nutrient enrichment on macroalgal coralligenous assemblages.

Effects of eutrophication on marine ecosystems have been widely studied, even if both the effects on deep subtidal rocky assemblages and response of different successional stages to nutrients impact are still not clear. In this context, the study aimed to evaluate the effects of nutrient enrichment on Mediterranean macroalgal assemblages associated with coralligenous habitat. A manipulative field experiment was carried out by supplying both mature and early successional stages of assemblages with nutrients. A total of 62 macroalgal species were identified. Multivariate and univariate analyses showed that the structure of both mature and early successional macroalgal assemblages of coralligenous significantly varied between areas treated with nutrients and not treated areas. Moreover, differences were stronger when macroalgal assemblages were in the early successional stage than in the mature one. Results highlighted the role played by nutrients in determining the structure of macroalgal coralligenous assemblages, furthermore suggesting possible synergetic effects with other kinds of disturbances.