Epilithic Bacterial Assemblages on Subtidal Rocky Reefs: Variation Among Alternative Habitats at Ambient and Enhanced Nutrient Levels.

Temperate rocky reefs often support mosaics of alternative habitats such as macroalgal forests, algal turfs and sea urchin barrens. Although the composition of epilithic microbial biofilms (EMBs) is recognized as a major determinant of macroalgal recruitment, their role in regulating the stability of alternative habitats on temperate rocky reefs remains unexplored. On shallow rocky reefs of the Island of Capraia (NW Mediterranean), we compared EMB structure among canopy stands formed by the fucoid Ericaria brachycarpa, algal turfs, and urchin barrens under ambient versus experimentally enhanced nutrient levels. The three habitats shared a core microbial community consisting of 21.6 and 25.3% of total ASVs under ambient and enhanced nutrient conditions, respectively. Although Gammaproteobacteria, Alphaproteobacteria and Flavobacteriia were the most abundant classes across habitats, multivariate analyses at the ASV level showed marked differences in EMB composition among habitats. Enhancing nutrient level had no significant effect on EMBs, although it increased their similarity between macroalgal canopy and turf habitats. At both ambient and enriched nutrient levels, ASVs mostly belonging to Proteobacteria and Bacteroidetes were more abundant in EMBs from macroalgal canopies than barrens. In contrast, ASVs belonging to the phylum of Proteobacteria and, in particular, to the families of Rhodobacteraceae and Flavobacteriaceae at ambient nutrient levels and of Rhodobacteraceae and Bacteriovoracaceae at enhanced nutrient levels were more abundant in turf than canopy habitats. Our results show that primary surfaces from alternative habitats that form mosaics on shallow rocky reefs in oligotrophic areas host distinct microbial communities that are, to some extent, resistant to moderate nutrient enhancement. Understanding the role of EMBs in generating reinforcing feedback under different nutrient loading regimes appears crucial to advance our understanding of the mechanisms underpinning the stability of habitats alternative to macroalgal forests as well as their role in regulating reverse shifts.

Canopy-forming macroalgae can adapt to marine heatwaves.

Seawater warming and marine heatwaves (MHWs) have a major role on the fragmentation and loss of coastal marine habitats. Understanding the resilience and potential for adaptation of marine habitat forming species to ocean warming becomes pivotal for predicting future changes, improving present conservation and restoration strategies. In this study, a thermo-tolerance experiment was conducted to investigate the physiological effects of short vs long MHWs occurring at different timing on recruits of Gongolaria barbata, a canopy-forming species widespread in the Mediterranean Sea. The recruits were collected from a population of the Marine Protected Area of Porto Cesareo (Apulia, Ionian Sea). Recruits length, PSII maximal photochemical efficiency (Fv/Fm), photosynthetic pigments content, concentrations of antioxidant compounds and total antioxidant activity (DPPH) were the response variables measured during the experiment. Univariate asymmetrical analyses highlighted that all physiological variables were significantly affected by both the duration and the timing of the thermal stress with the only exception of recruits length. The higher Fv/Fm ratio, chlorophylls and carotenoids content, and antioxidant compounds concentration in recruits exposed to long-term stress likely indicate an acclimation of thalli to the new environmental conditions and hence, an increased tolerance of G. barbata to thermal stress. Results also suggest that the mechanisms of adaptation activated in response to thermal stress did not affect the natural growth rate of recruits. Overall, this study supports the hypothesis that canopy-forming species can adapt to future climate conditions demonstrating a physiological acclimation to cope with MHWs, providing strong evidence that adaptation of marine species to thermal stress is more frequent than expected, this contributing to design tailored conservation and restoration strategies for marine coastal habitat.

The challenge of setting restoration targets for macroalgal forests under climate changes.

The process of site selection and spatial planning has received scarce attention in the scientific literature dealing with marine restoration, suggesting the need to better address how spatial planning tools could guide restoration interventions. In this study, for the first time, the consequences of adopting different restoration targets and criteria on spatial restoration prioritization have been assessed at a regional scale, including the consideration of climate changes. We applied the decision-support tool Marxan, widely used in systematic conservation planning on Mediterranean macroalgal forests. The loss of this habitat has been largely documented, with limited evidences of natural recovery. Spatial priorities were identified under six planning scenarios, considering three main restoration targets to reflect the objectives of the EU Biodiversity Strategy for 2030. Results show that the number of suitable sites for restoration is very limited at basin scale, and targets are only achieved when the recovery of 10% of regressing and extinct macroalgal forests is planned. Increasing targets translates into including unsuitable areas for restoration in Marxan solutions, amplifying the risk of ineffective interventions. Our analysis supports macroalgal forests restoration and provides guiding principles and criteria to strengthen the effectiveness of restoration actions across habitats. The constraints in finding suitable areas for restoration are discussed, and recommendations to guide planning to support future restoration interventions are also included.

Marine heatwaves drive recurrent mass mortalities in the Mediterranean Sea.

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. Overall, we show that increasing the resolution of empirical observation is critical to enhancing our ability to more effectively understand and manage the consequences of climate change.

An integrated assessment of the Good Environmental Status of Mediterranean Marine Protected Areas.

Local, regional and global targets have been set to halt marine biodiversity loss. Europe has set its own policy targets to achieve Good Environmental Status (GES) of marine ecosystems by implementing the Marine Strategy Framework Directive (MSFD) across member states. We combined an extensive dataset across five Mediterranean ecoregions including 26 Marine Protected Areas (MPAs), their reference unprotected areas, and a no-trawl case study. Our aim was to assess if MPAs reach GES, if their effects are local or can be detected at ecoregion level or up to a Mediterranean scale, and which are the ecosystem components driving GES achievement. This was undertaken by using the analytical tool NEAT (Nested Environmental status Assessment Tool), which allows an integrated assessment of the status of marine systems. We adopted an ecosystem approach by integrating data from several ecosystem components: the seagrass Posidonia oceanica, macroalgae, sea urchins and fish. Thresholds to define the GES were set by dedicated workshops and literature review. In the Western Mediterranean, most MPAs are in good/high status, with P. oceanica and fish driving this result within MPAs. However, GES is achieved only at a local level, and the Mediterranean Sea, as a whole, results in a moderate environmental status. Macroalgal forests are overall in bad condition, confirming their status at risk. The results are significantly affected by the assumption that discrete observations over small spatial scales are representative of the total extension investigated. This calls for large-scale, dedicated assessments to realistically detect environmental status changes under different conditions. Understanding MPAs effectiveness in reaching GES is crucial to assess their role as sentinel observatories of marine systems. MPAs and trawling bans can locally contribute to the attainment of GES and to the fulfillment of the MSFD objectives. Building confidence in setting thresholds between GES and non-GES, investing in long-term monitoring, increasing the spatial extent of sampling areas, rethinking and broadening the scope of complementary tools of protection (e.g., Natura 2000 Sites), are indicated as solutions to ameliorate the status of the basin.

A fast-moving target: achieving marine conservation goals under shifting climate and policies.

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.

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.

Linking patterns and processes across scales: the application of scale-transition theory to algal dynamics on rocky shores.

Understanding how species and environments respond to global anthropogenic disturbances is one of the greatest challenges for contemporary ecology. The ability to integrate modeling, correlative and experimental approaches within individual research programs will be key to address large-scale, long-term environmental problems. Scale-transition theory (STT) enables this level of integration, providing a powerful framework to link ecological patterns and processes across spatial and temporal scales. STT predicts the large-scale (e.g. regional) behavior of a system on the basis of nonlinear population models describing local (e.g. patch-scale) dynamics and the interaction between these nonlinearities and spatial variation in population abundance or environmental conditions. Here we use STT to predict the dynamics of turf-forming algae on rocky shores at Capraia Island, in the northwest Mediterranean. We developed a model of algal turf dynamics based on density-dependent growth that included the effects of local interactions with canopy algae. The model was parameterized with field data and used to scale up the dynamics of algal turfs from the plot scale (20×20 cm) to the island scale (tens of km). The interaction between nonlinear growth and spatial variance in cover of turfing algae emerged as a key term to translate the local dynamics up to the island scale. The model successfully predicted short-term and long-term mean values of turf cover estimated independently from a separate experiment. These results illustrate how STT can be used to identify the relevant mechanisms that drive large-scale changes in ecological communities. We argue that STT can contribute significantly to the connection between biomechanics and ecology, a synthesis that is at the core of the emerging field of ecomechanics.

Can we preserve and restore overlooked macroalgal forests?

Habitat degradation and loss are severely affecting macroalgal forests worldwide, and their successful mitigation depends on the identification of the drivers of loss and the implementation of effective conservation and restoration actions. We made an extensive literature review 1- to document the historical (1789-1999) and recent (2000-2020) occurrence of the genus Cystoseira, Ericaria and Gongolaria reported in the literature along the 8000 km of the coasts of Italy, 2- to assess their decline and patterns of extinction, 3- to ascertain the drivers responsible for these changes, 4- to highlight the existence of success stories in their conservation and natural recovery. In the last twenty years, overall information on the distribution of Cystoseira s.l. exponentially increased, although research focused almost exclusively on intertidal reefs. Despite the lack of systematic monitoring programs, the local extinction of 371 populations of 19 different species of Cystoseira s.l. was documented across several regions, since 2000. Coastal engineering and poor quality of waters due to urban, agricultural or industrial activities were often documented as leading causes of habitat loss. However, the drivers of extinction were actually unknown for the majority of the populations and cause-effects relationships are scarcely documented. Although the proportion of protected populations increased to 77.8%, Marine Protected Areas are unlikely to guarantee adequate conservation efficacy, possibly also for the widespread lack of management and monitoring plans dealing specifically with Cystoseira s.l. species, and few evidences of natural recovery were observed. Our review shows the dramatic lack of baseline information for macroalgal forests, highlighting the urgent need for the monitoring of less accessible habitats, the collection of long-term data to unveil drivers of loss, and an updated reporting about the conservation status of the species of interest to plan future interventions.

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 date mussel Lithophaga lithophaga: Biology, ecology and the multiple impacts of its illegal fishery.

The date mussel Lithophaga lithophaga is an edible endolithic bivalve, protected by the EU Habitats Directive and other international agreements, living inside carbonate rocks. Its illegal harvesting is carried by breaking the rocks where the bivalve grows. The impact has cascade consequences as it causes permanent changes in the substrate characteristics, the removal of benthic species, a shift from highly complex to structurally simplified habitats. As a result, the rich biodiversity of rocky reefs turns into a biological desert, named "barren". Along with the over exploitation of fish, this practice leads to the increase of sea urchin density and grazing pressure on habitats, hampering the resilience of the associated biodiversity and functions. This paper summarizes the information on date mussel biology, ecology, ecotoxicology, fishery and the legal framework regulating its protection. Evidence indicates that illegal harvesting is still operated and widespread along the Mediterranean and has huge costs in terms of loss of natural capital and ecosystem services, and in terms of active ecological restoration. Two case study areas (the Sorrento and Salento peninsulas) were selected to assess the economic costs of this practice. Tangible economic costs in terms of ecosystems services' loss are huge (from ca. 35,000 to more than 400,000 euros/year in 6.6 km of Sorrento and ca. 1.8 million euros/year along the 69 km of Salento). These costs are, on average, ca. 30 times lower than those of ecosystem restoration. Data mining from websites indicates that date mussels are presently commercialized in hundreds of restaurants in Greece, Balkan countries, Spain and Italy, favoured also by the lack of appropriate consumer information. This practice should be controlled and contrasted at local scale, enforced by national legislations, and implemented by transnational initiatives. Social campaigns are needed to increase public awareness of the serious consequences of date-mussel fishery and consumption.

Enhanced nutrient loading and herbivory do not depress the resilience of subtidal canopy forests in Mediterranean oligotrophic waters.

The interaction between top-down and bottom-up forces determines the recovery trajectory of macroalgal forests exposed to multiple stressors. In an oligotrophic system, we experimentally investigated how nutrient inputs affected the recovery of Cystoseira brachycarpa following physical disturbance of varying intensities, both inside forested areas and at the boundary with sea urchin barrens. Unexpectedly, Cystoseira forests were highly resilient to disturbance, as they were able to recover from any partial damage. In general, the addition of nutrients sped up the recovery of Cystoseira. Thus, only the total canopy removal, in combination with either low nutrient availability or intense grazing pressure, promoted the expansion of mat-forming algae or urchin barrens, respectively. Our study suggests that the effects of enhanced nutrient levels may vary according to the trophic characteristics of the waterbody, and hence, are likely to vary among regions of the Mediterranean basin.

Are we ready for scaling up restoration actions? An insight from Mediterranean macroalgal canopies.

Extensive loss of macroalgal forests advocates for large-scale restoration interventions, to compensate habitat degradation and recover the associated ecological functions and services. Yet, restoration attempts have generally been limited to small spatial extensions, with the principal aim of developing efficient restoration techniques. Here, the success of outplanting Cystoseira amentacea v. stricta germlings cultured in aquaria was experimentally explored at a scale of tens of kms, by means of a multifactorial experimental design. In the intertidal rocky shores of SE Italy, locations with a continuous distribution for hundreds of meters or with few thalli forming patches of few centimeters of C. amentacea canopy were selected. In each location, the effects of adult conspecifics and the exclusion of macrograzers (salema fish and sea urchins) on the survival of germlings were tested. We evaluated the most critical determinants of mortality for germlings, including the overlooked pressure of mesograzers (e.g. amphipods, small mollusks, polychaetes). Despite the high mortality observed during outplanting and early settlement stages, survival of C. amentacea germlings was consistently favored by the exclusion of macrograzers, while the presence of adult conspecifics had no effects. In addition, the cost analysis of the interventions showed the feasibility of the ex-situ method, representing an essential tool for preserving Cystoseira forests. Large scale restoration is possible but requires baseline information with an in-depth knowledge of the species ecology and of the areas to be restored, together with the development of specific cultivation protocols to make consistently efficient restoration interventions.

Effects of a glyphosate-based herbicide on Fucus virsoides (Fucales, Ochrophyta) photosynthetic efficiency.

Herbicides are increasingly recognised as sources of water pollution. Glyphosate-based herbicides (GBHs) are widely used because of their low cost and high effectiveness. By measuring the photosynthetic efficiency of Fucus virsoides fronds exposed to a GBH (Roundup® Power 2.0), we investigated the effect of a continuous exposure (6 days) and the potential of recovery after a short exposure (24 h). Both experiments were carried out combining GBH with and without nutrient enrichment, simulating a runoff event. A factorial experimental design allowed us to assess the potential of interactions between GBH and nutrients, which are likely to co-occur in coastal areas. Our results show deleterious effects of GBH at low concentration on F. virsoides, independently from the duration of exposure and the presence of nutrients.

Nutrient Loading Fosters Seagrass Productivity Under Ocean Acidification.

The effects of climate change are likely to be dependent on local settings. Nonetheless, the compounded effects of global and regional stressors remain poorly understood. Here, we used CO2 vents to assess how the effects of ocean acidification on the seagrass, Posidonia oceanica, and the associated epiphytic community can be modified by enhanced nutrient loading. P. oceanica at ambient and low pH sites was exposed to three nutrient levels for 16 months. The response of P. oceanica to experimental conditions was assessed by combining analyses of gene expression, plant growth, photosynthetic pigments and epiphyte loading. At low pH, nutrient addition fostered plant growth and the synthesis of photosynthetic pigments. Overexpression of nitrogen transporter genes following nutrient additions at low pH suggests enhanced nutrient uptake by the plant. In addition, enhanced nutrient levels reduced the expression of selected antioxidant genes in plants exposed to low pH and increased epiphyte cover at both ambient and low pH. Our results show that the effects of ocean acidification on P. oceanica depend upon local nutrient concentration. More generally, our findings suggest that taking into account local environmental settings will be crucial to advance our understanding of the effects of global stressors on marine systems.

Experimental Perturbations Modify the Performance of Early Warning Indicators of Regime Shift.

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.

Deterministic Factors Overwhelm Stochastic Environmental Fluctuations as Drivers of Jellyfish Outbreaks.

Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the future.

Aspects of benthic decapod diversity and distribution from rocky nearshore habitat at geographically widely dispersed sites.

Relationships of diversity, distribution and abundance of benthic decapods in intertidal and shallow subtidal waters to 10 m depth are explored based on data obtained using a standardized protocol of globally-distributed samples. Results indicate that decapod species richness overall is low within the nearshore, typically ranging from one to six taxa per site (mean = 4.5). Regionally the Gulf of Alaska decapod crustacean community structure was distinguishable by depth, multivariate analysis indicating increasing change with depth, where assemblages of the high and mid tide, low tide and 1 m, and 5 and 10 m strata formed three distinct groups. Univariate analysis showed species richness increasing from the high intertidal zone to 1 m subtidally, with distinct depth preferences among the 23 species. A similar depth trend but with peak richness at 5 m was observed when all global data were combined. Analysis of latitudinal trends, confined by data limitations, was equivocal on a global scale. While significant latitudinal differences existed in community structure among ecoregions, a semi-linear trend in changing community structure from the Arctic to lower latitudes did not hold when including tropical results. Among boreal regions the Canadian Atlantic was relatively species poor compared to the Gulf of Alaska, whereas the Caribbean and Sea of Japan appeared to be species hot spots. While species poor, samples from the Canadian Atlantic were the most diverse at the higher infraordinal level. Linking 11 environmental variables available for all sites to the best fit family-based biotic pattern showed a significant relationship, with the single best explanatory variable being the level of organic pollution and the best combination overall being organic pollution and primary productivity. While data limitations restrict conclusions in a global context, results are seen as a first-cut contribution useful in generating discussion and more in-depth work in the still poorly understood field of biodiversity distribution.

Spatial relationships between polychaete assemblages and environmental variables over broad geographical scales.

This study examined spatial relationships between rocky shore polychaete assemblages and environmental variables over broad geographical scales, using a database compiled within the Census of Marine Life NaGISA (Natural Geography In Shore Areas) research program. The database consisted of abundance measures of polychaetes classified at the genus and family levels for 74 and 93 sites, respectively, from nine geographic regions. We tested the general hypothesis that the set of environmental variables emerging as potentially important drivers of variation in polychaete assemblages depend on the spatial scale considered. Through Moran's eigenvector maps we indentified three submodels reflecting spatial relationships among sampling sites at intercontinental (>10,000 km), continental (1000-5000 km) and regional (20-500 km) scales. Using redundancy analysis we found that most environmental variables contributed to explain a large and significant proportion of variation of the intercontinental submodel both for genera and families (54% and 53%, respectively). A subset of these variables, organic pollution, inorganic pollution, primary productivity and nutrient contamination was also significantly related to spatial variation at the continental scale, explaining 25% and 32% of the variance at the genus and family levels, respectively. These variables should therefore be preferably considered when forecasting large-scale spatial patterns of polychaete assemblages in relation to ongoing or predicted changes in environmental conditions. None of the variables considered in this study were significantly related to the regional submodel.