title: The effects of an invasive seaweed on native communities vary along a gradient of land-based human impacts.
abstract: The difficulty in teasing apart the effects of biological invasions from those of other anthropogenic perturbations has hampered our understanding of the mechanisms underpinning the global biodiversity crisis. The recent elaboration of global-scale maps of cumulative human impacts provides a unique opportunity to assess how the impact of invaders varies among areas exposed to different anthropogenic activities. A recent meta-analysis has shown that the effects of invasive seaweeds on native biota tend to be more negative in relatively pristine than in human-impacted environments. Here, we tested this hypothesis through the experimental removal of the invasive green seaweed, Caulerpa cylindracea, from rocky reefs across the Mediterranean Sea. More specifically, we assessed which out of land-based and sea-based cumulative impact scores was a better predictor of the direction and magnitude of the effects of this seaweed on extant and recovering native assemblages. Approximately 15 months after the start of the experiment, the removal of C. cylindracea from extant assemblages enhanced the cover of canopy-forming macroalgae at relatively pristine sites. This did not, however, result in major changes in total cover or species richness of native assemblages. Preventing C. cylindracea re-invasion of cleared plots at pristine sites promoted the recovery of canopy-forming and encrusting macroalgae and hampered that of algal turfs, ultimately resulting in increased species richness. These effects weakened progressively with increasing levels of land-based human impacts and, indeed, shifted in sign at the upper end of the gradient investigated. Thus, at sites exposed to intense disturbance from land-based human activities, the removal of C. cylindracea fostered the cover of algal turfs and decreased that of encrusting algae, with no net effect on species richness. Our results suggests that competition from C. cylindracea is an important determinant of benthic assemblage diversity in pristine environments, but less so in species-poor assemblages found at sites exposed to intense disturbance from land-based human activities, where either adverse physical factors or lack of propagules may constrain the number of potential native colonizers. Implementing measures to reduce the establishment and spread of C. cylindracea in areas little impacted by land-based human activities should be considered a priority for preserving the biodiversity of Mediterranean shallow rocky reefs.
authors.affiliation: Department of Biology, University of Pisa , Pisa , Italy.
authors.forename: Fabio
authors.initials: F
authors.lastname: Bulleri
authors.affiliation: CNR-IAMC, Istituto per l'Ambiente Marino Costiero , Castellammare del Golfo , Italy.
authors.forename: Fabio
authors.initials: F
authors.lastname: Badalamenti
authors.affiliation: Ruđer Bošković Institute, Center for Marine Research, G. Paliaga , Rovinj , Croatia.
authors.forename: Ljiljana
authors.initials: L
authors.lastname: Iveša
authors.affiliation: CNR-IAMC, Istituto per l'Ambiente Marino Costiero , Castellammare del Golfo , Italy.
authors.forename: Barbara
authors.initials: B
authors.lastname: Mikac
authors.affiliation: CNR-IAMC, Istituto per l'Ambiente Marino Costiero , Castellammare del Golfo , Italy.
authors.forename: Luigi
authors.initials: L
authors.lastname: Musco
authors.affiliation: Ruđer Bošković Institute, Center for Marine Research, G. Paliaga , Rovinj , Croatia.
authors.forename: Andrej
authors.initials: A
authors.lastname: Jaklin
authors.affiliation: Department of Biology, University of Pisa , Pisa , Italy.
authors.forename: Alex
authors.initials: A
authors.lastname: Rattray
authors.affiliation: CNR-IAMC, Istituto per l'Ambiente Marino Costiero, Castellammare del Golfo, Italy; Stazione Zoologica Anton Dohrn, Napoli, Italy.
authors.forename: Tomás
authors.initials: T
authors.lastname: Vega Fernández
authors.affiliation: Department of Biology, University of Pisa , Pisa , Italy.
authors.forename: Lisandro
authors.initials: L
authors.lastname: Benedetti-Cecchi
keywords: Benthic community; Biological invasions; Caulerpa cylindracea; Cumulative human impacts; Habitat degradation; Marine biodiversity; Mediterranean Sea; Non-native species; Seaweeds; Temperate rocky reefs