Impact of a nuclear power station effluent on marine forests: A case study in SE Brazil and insights for global warming scenarios.

One of the main disturbances caused by coastal nuclear power plants is the discharge of thermal effluents capable of affecting a number of marine systems, including macroalgal forests that support key ecosystem services such as carbon uptake, fisheries increment and coastal protection. This study aimed at describing the long-term trend (1992-2022) in the abundance of Sargassum forests from sites located inside and outside areas affected by the thermal effluent discharged by the Brazilian Nuclear Power Station (BNPS) and at evaluating the relationship between Sargassum cover and seawater temperature. This information is interesting to provide insights on whether and how Sargassum populations would likely be affected by increasing temperature due to climate change. We detected a long-term decline in Sargassum cover inside, but not outside the area affected by the BNPS thermal plume. Mean summer surface seawater temperature above 30 °C was identified as an important factor driving the decline of Sargassum abundance. This study highlights the impact caused by decades of discharge of the BNPS thermal effluent on Sargassum forests, which leads to predict the likely disappearance of marine forests under a climate change scenario in other sites situated in warm temperate regions.

Variations in epilithic microbial biofilm composition and recruitment of a canopy-forming alga between pristine and urban rocky shores.

Brown algae of the genus Ericaria are habitat formers on Mediterranean rocky shores supporting marine biodiversity and ecosystem functioning. Their population decline has prompted attempts for restoration of threatened populations. Although epilithic microbial biofilms (EMBs) are determinant for macroalgal settlement, their role in regulating the recovery of populations through the recruitment of new thalli is yet to be explored. In this study, we assessed variations in microbial biofilms composition on the settlement of Ericaria amentacea at sites exposed to different human pressures. Artificial fouling surfaces were deployed in two areas at each of three study sites in the Ligurian Sea (Capraia Island, Secche della Meloria and the mainland coast of Livorno), to allow bacterial biofilm colonization. In the laboratory, zygotes of E. amentacea were released on these surfaces to evaluate the survival of germlings. The EMB's composition was assessed through DNA metabarcoding analysis, which revealed a difference between the EMB of Capraia Island and that of Livorno. Fouling surfaces from Capraia Island had higher rates of zygote settlement than the other two sites. This suggests that different environmental conditions can influence the EMB composition on substrata, possibly influencing algal settlement rate. Assessing the suitability of rocky substrata for E. amentacea settlement is crucial for successful restoration.

Distribution of a canopy-forming alga along the Western Atlantic Ocean under global warming: The importance of depth range.

Sargassum species are among the most important canopy-forming algae in the Western Atlantic Ocean (WAO), providing habitat for many species and contributing to carbon uptake. The future distribution of Sargassum and other canopy-forming algae has been modelled worldwide, indicating that their occurrence in many regions is threatened by increased seawater temperature. Surprisingly, despite the recognized variation in vertical distribution of macroalgae, these projections generally do not evaluate their results at different depth ranges. This study aimed to project the potential current and future distributions of the common and abundant benthic Sargassum natans in the WAO (from southern Argentina to eastern Canada), under RCP 4.5 and 8.5 climate change scenarios, through an ensemble SDM approach. Possible changes between present and future distributions were assessed within two depth ranges, namely areas up to 20 m and areas up to 100 m depth. Our models forecast different distributional trends for benthic S. natans depending on the depth range. Up to 100 m, suitable areas for the species will increase by 21% under RCP 4.5, and by 15% under RCP 8.5, when compared to the potential current distribution. On the contrary, up to 20 m, suitable areas for the species will decrease by 4% under RCP 4.5 and by 14% under RCP 8.5, when compared to the potential current distribution. Under the worst scenario, losses up to 20 m depth will affect approximately 45,000 km2 of coastal areas across several countries and regions of WAO, with likely negative consequences for the structure and dynamics of coastal ecosystems. These findings highlight the importance of considering different depth ranges when building and interpreting predictive models of the distribution of habitat-forming subtidal macroalgae under climate change.

The Fucales Index: A new tool for monitoring subtidal rocky habitats, and its application to an Atlantic bay subjected to nuclear power plant's effluents.

Ecological indices are useful tools for environmental managers to monitor and detect changes caused by natural or anthropogenic disturbances. Despite the fact that descriptors of Fucales species are often included in indices for evaluating changes in coastal marine habitats, there is no index based solely on Fucales abundance. This study proposes the Fucales Index (FI), based on four classes of abundance of a selected Fucales species. The ability of FI to detect changes in the abundance of Sargassum sp. was tested in a Brazilian bay that is subjected to the effluent plume from a nuclear power plant. FI was significantly different when comparing areas inside and outside of the plume area, and it increased with increasing distance from the source of the disturbance. These findings suggest that FI is a suitable tool for assessing the effect of an effluent plume and potentially of other disturbances on rocky shores hosting Fucales.

Global COVID-19 lockdown highlights humans as both threats and custodians of the environment.

The global lockdown to mitigate COVID-19 pandemic health risks has altered human interactions with nature. Here, we report immediate impacts of changes in human activities on wildlife and environmental threats during the early lockdown months of 2020, based on 877 qualitative reports and 332 quantitative assessments from 89 different studies. Hundreds of reports of unusual species observations from around the world suggest that animals quickly responded to the reductions in human presence. However, negative effects of lockdown on conservation also emerged, as confinement resulted in some park officials being unable to perform conservation, restoration and enforcement tasks, resulting in local increases in illegal activities such as hunting. Overall, there is a complex mixture of positive and negative effects of the pandemic lockdown on nature, all of which have the potential to lead to cascading responses which in turn impact wildlife and nature conservation. While the net effect of the lockdown will need to be assessed over years as data becomes available and persistent effects emerge, immediate responses were detected across the world. Thus, initial qualitative and quantitative data arising from this serendipitous global quasi-experimental perturbation highlights the dual role that humans play in threatening and protecting species and ecosystems. Pathways to favorably tilt this delicate balance include reducing impacts and increasing conservation effectiveness.

Resurvey of sea urchins and mussels at protected and harvested shores a decade after: A beyond-BACI approach.

Comparing temporal patterns of distribution and abundance of target organisms between protected and harvested shores is essential to assess the extant effectiveness of marine protected areas (MPAs) and whether it is maintained through time. By means of an adapted Beyond-BACI approach, we compared the short- and long-term patterns of variation in the abundance of the sea urchin Paracentrotus lividus and the mussel Mytilus galloprovincialis at a protected shore (within the Parque Litoral Norte MPA, Portugal) and at three adjacent shores subject to intense harvesting over a decadal interval. Despite the existence of the MPA for more than 30 years, we did not obtain clear evidence of its persistent or recent effectiveness on intertidal species of commercial interest. We suggest the need for refining management options along the northern Portuguese coast, possibly by better enforcing current regulations and reconsidering the present design of protection schemes. Moreover, the adopted analytical approach may represent a methodological reference for similar investigations in systems where the perturbation of interest (protection or disturbance) would not occur at a given time during the course of the study, but has been operating since before the first sampling occasion and maintained until subsequent surveys.

Sea urchin chronicles. The effect of oxygen super-saturation and marine polluted sediments from Bagnoli-Coroglio Bay on different life stages of the sea urchin Paracentrotus lividus.

In marinas and harbours, the accumulation of pollutants in sediments, combined with poor exchange of water with the open sea, poses a major environmental threat. The presence of photosynthetic organisms and the related oxygen production, however, may alleviate the negative effects of environmental contamination on heterotrophic organisms, enhancing their physiological defences. Furthermore, possible transgenerational buffer effects may increase the ability of natural populations to face environmental stress. Here we tested the occurrence of transgenerational effects on larvae of the sea urchin Paracentrotus lividus, whose parents were exposed, during the gametogenesis, to contaminated sediments subject to two temporal patterns of water re-suspension events and normal- (90%) vs. super-saturated (200%) levels of O2. The study site was Bagnoli-Coroglio (Gulf of Naples, southern Tyrrhenian Sea), a historically polluted brownfield and Site of National Interest for which environmental restoration options are currently under exploration. Larvae from different adult populations were significantly, although not linearly, affected by the interaction of all factors to which parents were exposed, at both 24h and 48h post fertilization. Specifically, the exposure of larvae to elutriates from contaminated sediments determined a developmental delay, a reduction in size and an increased percentage of abnormalities in all larval populations independently of their parental exposure. On the contrary, larvae from parents exposed to contaminated sediments, when reared in clean filtered sea water, succeeded in developing until the echinopluteus stage after 48h, with size and abundance comparable to those of larvae from control parents. Pre-exposure of parents to contaminated sediments did not successfully buffer the negative effects of elutriates on their offspring, and no positive effects of 'super-saturated' levels of O2 in response to contaminants were observed, suggesting that the Bagnoli-Coroglio area is currently not suitable for the re-stocking or re-introduction of this species.

Oxygen supersaturation mitigates the impact of the regime of contaminated sediment reworking on sea urchin fertilization process.

Dismissed industrial plants with chronic environmental contamination globally affect all levels of biological organization in concert with other natural and anthropogenic perturbations. Assessing the impact of such perturbations and finding effective ways to mitigate them have clear ecological and societal implications. Through indoor manipulative experiments, we assessed here the effects of the temporal regime of reworking of contaminated sediment from the Bagnoli-Coroglio brownfield (Tyrrhenian Sea, Italy) on the fertilization process in Paracentrotus lividus. Adult sea urchins were kept for one month in tanks containing contaminated sediment that was re-suspended according to two temporal patterns of water turbulence differing in the time intervals between consecutive events of agitation (mimicking the storms naturally occurring in the study area) in seawater with natural vs. supersaturated oxygenation levels. At the end of the treatment, gametes were collected and used to test the hypothesis that the regime of contaminated sediment reworking negatively, but reversibly, affects morphological and physiological traits of the fertilized eggs. We found that aggregated events of sediment re-suspension had profound negative effects on gamete interactions and Ca2+ signaling at fertilization. The same experimental condition also inflicted marked ultrastructural changes in eggs. Importantly, however, such detrimental effects were inhibited by increased oxygenation. By contrast, the regime of sediment re-working with a longer interval between consecutive turbulent events had only marginal effects. Thus, the current and predicted changes of climate-related disturbance appear to modulate the biological effects of chronic contamination in post-industrial areas, suggesting that environmental rehabilitation via restoration of habitat-forming primary producers such as seagrasses or algal canopies could alleviate the pollutants' effects on resident biota.

An indoor study of the combined effect of industrial pollution and turbulence events on the gut environment in a marine invertebrate.

Natural storms are able to determine reworking of seabed up to considerable depths and favour suspension of sediment-associated chemicals. Yet, a direct link between exposure to resuspended contaminants and the biological effects on marine organisms have to be fully established. We exposed adults of a suspension feeder, the ascidian Ciona robusta, to polluted sediment (e.g., containing mixtures of polycyclic aromatic hydrocarbons and heavy metals) from the industrial area of Bagnoli-Coroglio under two temporal patterns ('aggregated' vs. 'spaced') of turbulence events. Then, we assessed the impact of resuspended pollutants on the ascidian gut environment via four broad categories: oxidative stress, innate immunity, host-microbiota interactions, and epithelium. An early oxidative stress response was seen after a week of exposure to static sediment. Instead, water turbulence had no effect on the antioxidant defence. The first episode of turbulent suspension induced a minimal pro-inflammatory response in the 'spaced' pattern. Mucus overproduction and a complete occlusion of the crypt lumen were found following sediment reworking. This study suggests a protective response of the gut environment in marine invertebrates exposed to environmental extremes, leading to increased susceptibility to disease and to concerns on the combined effects of chronic environmental contamination and acute disturbance events possibly associated with climate change.

Morphological and molecular responses of the sea urchin Paracentrotus lividus to highly contaminated marine sediments: The case study of Bagnoli-Coroglio brownfield (Mediterranean Sea).

Marine sediments store complex mixtures of compounds, including heavy metals, organotins and a large array of other contaminants. Sediment quality monitoring, characterization and management are priorities, due to potential impacts of the above compounds on coastal waters and their biota, especially in cases of pollutants released during dredging activities. Harbours and marinas, as well as estuaries and bays, where limited exchanges of water occurr, the accumulation of toxic compounds poses major concerns for human and environmental health. Here we report the effects of highly contaminated sediments from the site of national interest Bagnoli-Coroglio (Tyrrhenian Sea, Western Mediterranean) on the sea urchin Paracentrotus lividus, considered a good model for ecotoxicological studies. Adult sea urchins were reared one month in aquaria in the presence of contaminated sediment that was experimentally subject to different patterns of re-suspension events (mimicking the effect of natural storms occurring in the field), crossed with O2 enrichment versus natural gas exchanges in the water. The development of embryos deriving from adult urchins exposed to such experimental conditions was followed until the pluteus stage, checking the power of contaminated sediment to induce morphological malformations and its eventual buffering by high oxygenation. Real-Time qPCR analysis revealed that the expression of several genes (among the fifty analyzed, involved in different functional processes) was targeted by contaminated sediments more than those exposed in oxygen-enriched condition. Our findings have biological and ecological relevance in terms of assessing the actual impact on local organisms of chronic environmental contamination by heavy metals and polycyclic aromatic hydrocarbons affecting the Bagnoli-Coroglio area, and of exploring enhanced sediment and water oxygenation as a promising tool to mitigate the effects of contamination in future environmental restoration actions.

Recovery ability of lowshore sessile assemblages in a highly contaminated post-industrial area.

The inheritance of environmental contamination left by abandoned industrial plants is widespread globally. Here we compared the patterns of recovery of lowshore algal and invertebrate assemblages between the post-industrial site of Bagnoli-Coroglio and four reference sites distributed along the coast in the Gulf of Naples, southern Tyrrhenian Sea. The structure of whole assemblages, richness of taxa and abundance of individual taxa were followed during one year since an event of experimental disturbance consisting in the removal of all erect organisms from the rocky substrate. Our main findings suggest that the examined benthic assemblages recovered effectively and quickly after a pulse disturbance and, contrarily to initial expectations, that this ability was comparable between the post-industrial site and the reference sites. This result is discussed in terms of several plausible processes and mechanisms, including the general capability of intertidal organisms to recover from physical disturbance, the potential high level of environmental stress affecting the reference sites too, the chance that the most intense impacts of contamination remained restricted to the sediments of the post-industrial site without propagating to adjacent rocky habitats, and the large natural variability of reference sites that may have masked weak effects of the historical contamination. Irrespective of the actual causes, we emphasize the need for including natural variability of the examined system in any future restoration interventions, to guarantee representation of the range of variation of target organisms and of their underlying processes, and to avoid confounding the intended post-industrial impact with the effects of other natural and anthropogenic processes.

Short-term variation of abundance of the purple sea urchin, Paracentrotus lividus (Lamarck, 1816), subject to harvesting in northern Portugal.

Paracentrotus lividus is a common and intensely harvested sea urchin at several European locations, including the Mediterranean Sea and the Atlantic coast of the Iberian Peninsula. The increasing human pressure on this resource due to the growing demand and market value of sea urchin gonads as seafood raises concerns on the ecological sustainability of present fisheries, which are showing a technological improvement and an expansion towards previously non-harvested areas. We examined the abundance of P. lividus of both commercial and non-commercial size before, during and after the harvesting season (from October to April) in the rocky shallow subtidal habitat along the northern Portuguese coast. The abundance of commercial (≥50 mm in test diameter) P. lividus individuals increased in the harvesting season, but drastically dropped by about 90% in the after-harvesting period. Such a pattern was consistent among three rocky shores spanning about 65 km of coast. The multivariate population structure and most size classes of non-commercial sea urchins did not differ depending on the period. The only exception was Class 4 (test diameter between 30 and 40 mm), which was more abundant in the harvesting than in the before- and, further, the after-harvesting period, but only at one shore. Very small (Class 1, test diameter below 10 mm) urchins were never found. The present findings suggest that human harvesting may cause considerable reductions in the abundance of target P. lividus, but that such an effect would not be evident concomitantly with harvesting, but in the subsequent period. Even if just under a precautionary principle, protection strategies focused on sea urchin populations and the harvesting period are advisable to contribute to maintain a sustainable local fishery of P. lividus populations that are likely to be negatively affected also by other natural and anthropogenic perturbations.

Reducing the data-deficiency of threatened European habitats: Spatial variation of sabellariid worm reefs and associated fauna in the Sicily Channel, Mediterranean Sea.

Biogenic reefs, such as those produced by tube-dwelling polychaetes of the genus Sabellaria, are valuable marine habitats which are a focus of protection according to European legislation. The achievement of this goal is potentially hindered by the lack of essential empirical data, especially in the Mediterranean Sea. This study addresses some of the current knowledge gaps by quantifying and comparing multi-scale patterns of abundance and distribution of two habitat-forming species (Sabellaria alveolata and S. spinulosa) and their associated fauna along 190 km of coast on the Italian side of the Sicily Channel. While the abundance of the two sabellariids and the total number of associated taxa did not differ at any of the examined scales (from tens of centimetres to tens-100 of kilometres), the structure (composition in terms of both the identity and the relative abundance of constituting taxa) of the associated fauna and the abundance of several taxa (the polychaetes Eulalia ornata, Syllis pulvinata, S. garciai, Nereis splendida and Arabella iricolor, and the amphipods Apolochus neapolitanus, Tethylembos viguieri and Caprella acanthifera) varied among locations established ∼50-100 km apart. Syllis pulvinata also showed significant variation between sites (hundreds of metres apart), analogously to the other syllid polychaetes S. armillaris and S. gracilis, the nereidid polychaete Nereis rava, and the amphipod Gammaropsis ulrici. The largest variance of S. spinulosa, of the structure of the whole associated fauna and of 56% of taxa analysed individually occurred at the scale of replicates (metres apart), while that of the dominant bio-constructor S. alveolata and of 25% of taxa occurred at the scale of sites. The remaining 19% and the total richness of taxa showed the largest variance at the scale of locations. Present findings contribute to meet a crucial requirement of any future effective protection strategy, i.e., identifying relevant scales of variation to be included in protection schemes aiming at preserving representative samples not only of target habitats and organisms, but also of the processes driving such variability.

Symbiont dynamics during thermal acclimation using cnidarian-dinoflagellate model holobionts.

Warming oceans menace reef ecosystems by disrupting symbiosis between cnidarians and Symbiodinium zooxanthellae, thus triggering bleach episodes. Temperature fluctuations promote adjustments in physiological variables and symbiont composition, which can cause stress responses, but can also yield adaptation if fitter host-symbiont homeostasis are achieved. To understand such processes manipulative studies are required, but many reef-building cnidarians pose limitations to experimental prospects. We exposed Exaiptasia anemones to Gradual Thermal Stress (GTS) and Heat Shock (HS) exposures and monitored chlorophyll and symbiont dynamics to test the phenotypic plasticity of these photosynthetic holobionts. GTS enhanced chlorophyll concentrations and decreased Symbiodinium proliferation. A recovery period after GTS returned chlorophyll to lower concentrations and symbiont divisions to higher rates. HS triggered a stress response characterized by intense symbiont declines through degradation and expulsion, algal compensatory proliferation, and chlorophyll accumulation. Anemones pre-exposed to GTS displayed more acute signs of symbiont paucity after HS, demonstrating that recurrent stress does not always induce bleaching-resistance. Our study is the first documenting Symbiodinium C and D, along with the predominant Clade B1 in Exaiptasia anemones. C subclades found in outdoor specimens faded under laboratory exposures. Clade D emerged after HS treatments, and especially after GTS pre-exposure. This highlights the thermotolerance of D subclades found in E. pallida and shows that bleaching-recovery can involve shifts of background symbiont phylotypes. This study enlightens the capability of Exaiptasia anemones to acclimate to gradually increased temperatures, and explores into how thermal history influences in subsequent stress tolerance in symbiotic cnidarians.

Nowhere safe? Exploring the influence of urbanization across mainland and insular seashores in continental Portugal and the Azorean Archipelago.

Differences in the structure and functioning of intensively urbanized vs. less human-affected systems are reported, but such evidence is available for a much larger extent in terrestrial than in marine systems. We examined the hypotheses that (i) urbanization was associated to different patterns of variation of intertidal assemblages between urban and extra-urban environments; (ii) such patterns were consistent across mainland and insular systems, spatial scales from 10scm to 100skm, and a three months period. Several trends emerged: (i) a more homogeneous distribution of most algal groups in the urban compared to the extra-urban condition and the opposite pattern of most invertebrates; (ii) smaller/larger variances of most organisms where these were, respectively, less/more abundant; (iii) largest variability of most response variables at small scale; (iv) no facilitation of invasive species by urbanization and larger cover of canopy-forming algae in the insular extra-urban condition. Present findings confirm the acknowledged notion that future management strategies will require to include representative assemblages and their relevant scales of variation associated to urbanization gradients on both the mainland and the islands.

Devil's tongue weed (Grateloupia turuturu Yamada) in northern Portugal: Passenger or driver of change in native biodiversity?

Understanding the mechanisms underlying biological invasions is essential to separate their actual ecological effects from those of other human disturbances. This study examined experimentally whether the non-native red seaweed Grateloupia turuturu is an opportunistic species taking advantage of degraded local conditions (passenger model), or the primary driver of changes in the structure of benthic assemblages (driver model). In a first experiment, traits of G. turuturu likely associated to its invasion success were compared between unmanipulated controls and treatments subjected to the removal of canopy-forming macroalgae to test for the passenger model. In a second experiment, rock pool assemblages where G. turuturu was selectively removed were compared with initially similar control (unmanipulated) assemblages to test for the driver model. Over a period of four months, G. turuturu showed larger cover, higher number of individuals and longer fronds in the canopy-removed compared to the control pools, while no significant differences were detected between assemblages where G. turuturu was continuously removed and those where invasion was allowed to occur. Present findings suggest that G. turuturu would rely on disturbances removing potential native competitors to spread into the recipient habitat, rather than being the main driver of ecological alterations.

Effects of subtle pollution at different levels of biological organisation on species-rich assemblages.

We investigated effects of subtle nutrient enrichment and metal pollution on different levels of biological organization (i.e. whole assemblage, population and individual) of species-rich assemblages. We used rockpools as model system, applying a multi-factorial sampling design to test hypotheses on differences between disturbed and reference locations. Results indicated that disturbed and reference locations supported similar assemblages, as well as individual fitness-related life-traits were ineffective to discriminate between the two conditions. In contrast, assemblages responded to pollution through a reduction of the abundance of sensitive species and a proliferation of tolerant species, although these alterations were detectable only once the influence of dominant taxa was down-weighed by data transformation. Present findings suggest that, contrarily to individual level variables, assemblage structure after data transformation and patterns of distribution and abundance of differently sensitive taxa would be a powerful tool to detect effects of subtle pollution on species-rich assemblages.

Reddened seascapes: experimentally induced shifts in 1/f spectra of spatial variability in rocky intertidal assemblages.

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.

Patterns of variation of intertidal species of commercial interest in the Parque Litoral Norte (north Portugal) MPA: comparison with three reference shores.

Marine Protected Areas (MPAs) are world-wide established with the aim of conserving biodiversity and preventing overexploitation of marine organisms. Evaluating the effectiveness of MPAs is needed in order to support and implement their management, but it is complicated by the large natural variability in space and time of distribution and abundance of natural populations. Here, we tested the hypothesis that patterns of total abundance and size-frequency distribution of two intensively harvested intertidal species (the sea urchin Paracentrotus lividus and the mussel Mytilus galloprovincialis) differed between a protected and three reference shores along the rocky coast of north Portugal. Response variables were in terms of mean values and measures of variance at different spatial scales (from centimetres to metres) and over time (along a period of about 12 months). A further comparison involved the estimation of the reproductive potential of sea urchins, quantified as variations of Gonad Index (GI = gonad dry weight/body dry weight × 100) at the scale of shore. Results did not generally support a predictable direct effect of protection, as the total abundance and the abundance of larger individuals of both species and GI did not differ between the MPA and reference shores. However, a considerable temporal and spatial variability at smaller scales was detected for several response variables. Such findings have implications for management of MPAs, highlighting the need for sampling designs properly replicated in space and time, in order to examine their effectiveness, and for considering spatial and temporal heterogeneity of target populations and driving processes as a criterion for their implementation and design.

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.