Abstract. 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.

Climate change is causing an increase in the frequency and intensity of marine heatwaves (MHWs) and mass mortality events (MMEs) of marine organisms are one of their main ecological impacts. Here, we show that during the 2015-2019 period, the Mediterranean Sea has experienced exceptional thermal conditions resulting in the onset of five consecutive years of widespread MMEs across the basin. These MMEs affected thousands of kilometers of coastline from the surface to 45 m, across a range of marine habitats and taxa (50 taxa across 8 phyla). Significant relationships were found between the incidence of MMEs and the heat exposure associated with MHWs observed both at the surface and across depths. Our findings reveal that the Mediterranean Sea is experiencing an acceleration of the ecological impacts of MHWs which poses an unprecedented threat to its ecosystems' health and functioning.

Ecosystems may shift abruptly between alternative states in response to environmental perturbations. Early warning indicators have been proposed to anticipate such regime shifts, but experimental field tests of their validity are rare. We exposed rocky intertidal algal canopies to a gradient of press perturbations and recorded the response of associated assemblages over 7 years. Reduced cover and biomass of algal canopies promoted the invasion of algal turfs, driving understory assemblages toward collapse upon total canopy removal. A dynamic model indicated the existence of a critical threshold separating the canopy- and turf-dominated states. We evaluated common indicators of regime shift as the system approached the threshold, including autocorrelation, SD, and skewness. These indicators captured changes in understory cover due to colonization of algal turfs. All indicators increased significantly as the system approached the critical threshold, in agreement with theoretical predictions. The performance of indicators changed when we superimposed a pulse disturbance on the press perturbation that amplified environmental noise. This treatment caused several experimental units to switch repeatedly between the canopy- and the turf-dominated state, resulting in a significant increase in overall variance of understory cover, a negligible effect on skewness and no effect on autocorrelation. Power analysis indicated that autocorrelation and SD were better suited at anticipating a regime shift under mild and strong fluctuations of the state variable, respectively. Our results suggest that regime shifts may be anticipated under a broad range of fluctuating conditions using the appropriate indicator.

Predicting community susceptibility to invasion has become a priority for preserving biodiversity. We tested the hypothesis that the occurrence and abundance of the seaweed Caulerpa racemosa in the north-western (NW) Mediterranean would increase with increasing levels of human disturbance. Data from a survey encompassing areas subjected to different human influences (i.e. from urbanized to protected areas) were fitted by means of generalized linear mixed models, including descriptors of habitats and communities. The incidence of occurrence of C. racemosa was greater on urban than extra-urban or protected reefs, along the coast of Tuscany and NW Sardinia, respectively. Within the Marine Protected Area of Capraia Island (Tuscan Archipelago), the probability of detecting C. racemosa did not vary according to the degree of protection (partial versus total). Human influence was, however, a poor predictor of the seaweed cover. At the seascape level, C. racemosa was more widely spread within degraded (i.e. Posidonia oceanica dead matte or algal turfs) than in better preserved habitats (i.e. canopy-forming macroalgae or P. oceanica seagrass meadows). At a smaller spatial scale, the presence of the seaweed was positively correlated to the diversity of macroalgae and negatively to that of sessile invertebrates. These results suggest that C. racemosa can take advantage of habitat degradation. Thus, predicting invasion scenarios requires a thorough knowledge of ecosystem structure, at a hierarchy of levels of biological organization (from the landscape to the assemblage) and detailed information on the nature and intensity of sources of disturbance and spatial scales at which they operate.

In the Anthropocene, marine ecosystems are rapidly shifting to new ecological states. Achieving effective conservation of marine biodiversity has become a fast‐moving target because of both global climate change and continuous shifts in marine policies. How prepared are we to deal with this crisis? We examined EU Member States Programs of Measures designed for the implementation of EU marine environmental policies, as well as recent European Marine Spatial Plans, and discovered that climate change is rarely considered operationally. Further, our analysis revealed that monitoring programs in marine protected areas are often insufficient to clearly distinguish between impacts of local and global stressors. Finally, we suggest that while the novel global Blue Growth approach may jeopardize previous marine conservation efforts, it can also provide new conservation opportunities. Adaptive management is the way forward (e.g., preserving ecosystem functions in climate change hotspots, and identifying and targeting climate refugia areas for protection) using Marine Spatial Planning as a framework for action, especially given the push for Blue Growth.

Ecological tests of 1/f-noise models have advanced our understanding of how environmental fluctuations affect population abundance and species distributions. Most empirical studies have been conducted under controlled laboratory conditions and have focused on individual drivers. We present the results of a four-year field experiment in which canopy presence/absence and the availability of primary space were manipulated as red-noise and white-noise spatial processes, respectively, to evaluate their separate and compounded effects on algal turf distribution in a rocky intertidal community. Algal turfs closely tracked spatial variation in canopy distribution, displaying a reddened spectrum of spatial variation. Surprisingly, white-noise clearings also induced a red-shift in turf distribution, a pattern that was related to a nonlinear relation between gap size and turf colonization. The two disturbances interacted antagonistically, dampening the red-shift of turf distribution. Our results provide evidence of experimentally induced shifts in the spectrum of a spatial variable under natural environmental conditions.

Despite the great interest for the role played by resident assemblages in regulating biological invasions, few studies have assessed how these can influence the spread of exotic species that have successfully established or have included more than one trophic level. On shallow rocky reefs, we assessed how the effects of different benthic assemblages on the spread of an invasive alga, Caulerpa racemosa, are influenced by alterations in the density and species composition of the resident sea urchin assemblage. In order to simulate herbivore species loss scenarios, assemblages dominated by different morphological groups of algae (i.e. turfs or encrusting corallines) or experimentally cleared plots (i.e. bare rock) were exposed to grazing by different combinations of species (Arbacia lixula and Paracentrotus lividus) and densities (natural, -50% and -100% of natural densities) of urchins. Algal turfs and encrusting corallines generally facilitated C. racemosa. Manipulating urchins assemblages did not affect the cover and density of fronds of C. racemosa. In contrast, halving the density of P. lividus favoured the penetration of stolons of C. racemosa, consistently among algal assemblages. Other effects of urchins varied among algal assemblages, indicating interactions between trophic levels. In algal turfs, the total removal of urchins caused a decrease in the penetration of stolons, while it enhanced the length of fronds, indicating a shift in the growth form of this clonal plant and, ultimately, a depression of its spreading ability. In bare or encrusting corallines dominated surfaces, the removal of urchins had positive effects on the penetration of stolons and on the length of fronds of C. racemosa, irrespective of one or two species being manipulated and for the intensity of their removal. Our results show that tradeoffs between negative and positive effects of herbivores, varying according to relative densities of species and to the direction and strength of the effects of resident plant assemblages, can influence local rates of spread of C. racemosa. Thus, not only facilitation of exotics by natives can be key in enhancing the spread of exotic species, but it can occur between organisms at different trophic levels. Finally, our findings have important implications for the management of C. racemosa, in view of the progressive domination of shallow rocky reefs by algal turfs

Compounded effects of climate change and local human activities are threatening marine biodiversity worldwide. At a regional scale (10s to 100s km), comparisons among areas characterized by the prevalence of different human activities provide an insight into the effects of anthropogenic disturbances at multiple levels of ecological organization (i.e. from landscapes to assemblages). At the landscape scale (1000s m), we hypothesized that patchiness in habitat distribution and proportion of degraded assemblages would increase with increasing levels of disturbance, as a result of the decline of habitat-forming species. In addition, we hypothesized that prevailing human influences would affect the structure and variability of rocky benthic assemblages at smaller spatial scales (10s cm to 10s m). An extensive survey encompassing areas subjected to different human influences (i.e. from urbanized to protected areas) was carried out along the coasts of Tuscany (NW Mediterranean Sea). Seagrass beds and macroalgal canopy stands were the dominant habitats in relatively pristine areas, while macroalgal turfs and dead rhizomes of Posidonia oceanica were the most extended habitats in urbanized areas. In general, habitat fragmentation did not vary among areas subjected to different human influences. At a smaller scale (10s cm to 10s m), urbanization favored dominance by opportunistic species and promoted biotic homogenization. Our study shows that regional variations in the composition of landscapes and assemblages can be predicted on the basis of prevailing human activities. Our results also suggest that variations in landscape composition could be an effective descriptor of the effects of multiple human stressors in marine environments.