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Although many studies predict extensive future biodiversity loss and redistribution in the terrestrial realm, future changes in marine biodiversity remain relatively unexplored. In this work, we model global shifts in one of the most important marine functional groups-ecosystem-structuring macrophytes-and predict substantial end-of-century change. By modelling the future distribution of 207 brown macroalgae and seagrass species at high temporal and spatial resolution under different climate-change projections, we estimate that by 2100, local macrophyte diversity will decline by 3-4% on average, with 17 to 22% of localities losing at least 10% of their macrophyte species. The current range of macrophytes will be eroded by 5-6%, and highly suitable macrophyte habitat will be substantially reduced globally (78-96%). Global macrophyte habitat will shift among marine regions, with a high potential for expansion in polar regions., (© 2024. The Author(s).)

Determining the proximity of ecosystems to tipping points is a critical yet complex task, heightened by the growing severity of climate change and local anthropogenic stressors on ecosystem integrity. Spatial Early Warning Signals (EWS) have been recognized for their potential in preemptively signaling regime shifts to degraded states, but their performance in natural systems remains uncertain. In this study, we investigated the performance of 'recovery length' - the spatial extent of recovery from a perturbation - and spatial EWS as early warnings of regime shifts in Posidonia oceanica meadows. Our experimental approach involved progressively thinning the P. oceanica canopy, from 0 to 100%, at the edge of a dead-matte area - a structure formed by dead P. oceanica rhizomes and colonized by algal turfs - to promote the propagation of algal turfs. We calculated recovery length as the distance from the dead-matte edge to the point where algal turfs colonized the canopy-thinned region. Our results showed a linear increase in recovery length with canopy thinning, successfully anticipated the degradation of P. oceanica. While spatial skewness decline with increased canopy degradation, other spatial EWS, such as Moran correlation at lag-1, low-frequency spatial spectra, and spatial variance, were ineffective in signaling this degradation. These findings underscore the potential of recovery length as a reliable early warning indicator of regime shifts in marine coastal ecosystems., Competing Interests: Declaration of competing interest The authours declare that they have no conflict that could influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

Understanding how synchronous species fluctuations affect community stability is a main research topic in ecology. Yet experimental studies evaluating how changes in disturbance regimes affect the synchrony and stability of populations and communities remain rare. We hypothesized that spatially heterogeneous disturbances of moderate intensity would promote metacommunity stability by decreasing the spatial synchrony of species fluctuations. To test this hypothesis, we exposed rocky shore communities of algae and invertebrates to homogeneous and gradient-like spatial patterns of disturbance at two levels of intensity for 4 years and used synchrony networks to characterize community responses to these disturbances. The gradient-like disturbance at low intensity enhanced spatial β diversity compared to the other treatments and produced the most heterogeneous and least synchronized network, which was also the most stable in terms of population and community fluctuations. In contrast, homogeneous disturbance destabilized the community, enhancing spatial synchronization. Intense disturbances always reduced spatial β diversity, indicating that strong perturbations could destabilize communities via biotic homogenization regardless of their spatial structure. Our findings corroborated theoretical predictions, emphasizing the importance of spatially heterogeneous disturbances in promoting stability by amplifying asynchronous spatial and temporal fluctuations in population and community abundance. In contrast to other networks, synchrony networks are vulnerable to the removal of most peripheral nodes, which are less synchronized, but may contribute more to stability than other nodes by dampening large fluctuations in species abundance. Our findings suggest that climate change and direct anthropogenic disturbance can compromise the stability of ecological communities through combined effects on diversity and synchrony, as well as further affecting ecosystems through habitat loss., (© 2024 The Ecological Society of America.)

As β-diversity can be seen as a proxy of ecological connections among species assemblages, modeling the decay of similarity in species composition at increasing distance may help elucidate spatial patterns of connectivity and local- to large-scale processes driving community assembly within a marine region. This, in turn, may provide invaluable information for setting ecologically coherent networks of marine protected areas (MPAs) in which protected communities are potentially interrelated and can mutually sustain against environmental perturbations. However, field studies investigating changes in β-diversity patterns at a range of spatial scales and in relation to disturbance are scant, limiting our understanding of how spatial ecological connections among marine communities may affect their recovery dynamics. We carried out a manipulative experiment simulating a strong physical disturbance on subtidal rocky reefs at several locations spanning >1000 km of coast in the Adriatic Sea (Mediterranean Sea) and compared β-diversity patterns and decay of similarity with distance and time by current transport between undisturbed and experimentally disturbed macrobenthic assemblages to shed light on connectivity processes and scales involved in recovery. In contrast to the expectation that very local-scale processes, such as vegetative regrowth and larval supply from neighboring undisturbed assemblages, might be the major determinants of recovery in disturbed patches, we found that connectivity mediated by currents at larger spatial scales strongly contributed to shape community reassembly after disturbance. Across our study sites in the Adriatic Sea, β-diversity patterns suggested that additional protected sites that matched hotspots of propagule exchange could increase the complementarity and strengthen the ecological connectivity throughout the MPA network. More generally, conditional to habitat distribution and selection of sites of high conservation priority (e.g., biodiversity hotspots), setting network internode distance within 100-150 km, along with sizing no-take zones to cover at least 5 km of coast, would help enhance the potential connectivity of Mediterranean subtidal rocky reef assemblages from local to large scale. These results can help improve conservation planning to achieve the goals of promoting ecological connectivity within MPA networks and enhancing their effectiveness in protecting marine communities against rapidly increasing natural and anthropogenic disturbances., (© 2023 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Coastal environments experience both natural and anthropogenic inputs of nitrogen (N) and phosphorus (P). Agricultural fertilisers, organic run-offs, and edaphic characteristics of coastal environments may generate mosaics of nutrient concentrations that ultimately influence the coastal primary productivity. Here, we experimentally assessed the effects of repeated pulses of N and P on multiple components of ecological stability (sensitivity, resilience, temporal stability and recovery) of phototrophic rocky intertidal biofilm. We performed a repeated-pulses factorial experiment crossing increasing N and P concentrations chosen to reflect a range of nutrient enrichment conditions, from oligotrophic to eutrophic. N and P, regardless of concentration or whether they occurred in isolation or combination, enhanced biofilm's sensitivity (increased biomass or physiological performance compared to controls) without altering resilience. Our experiment illustrates how the stability of an essential coastal primary producer responds to increasing N and P supply levels. Furthermore, notwithstanding the importance of decomposing the multiple dimensions of stability, the transitory increase of the sole sensitivity indicated that rocky shore biofilm is robust against a wide range of nutrient enrichment., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Network theory offers innovative tools to explore the complex ecological mechanisms regulating species associations and interactions. Although interest in ecological networks has grown steadily during the last two decades, the application of network approaches has been unequally distributed across different study systems: while some kinds of interactions (e.g. plant-pollinator and host-parasite) have been extensively investigated, others remain relatively unexplored. Among the latter, aquatic macrophyte-animal associations in coastal environments have been largely neglected, despite their major role in littoral ecosystems. The ubiquity of macrophyte systems, their accessibility and multi-faceted ecological, economical and societal importance make macrophyte-animal systems an ideal subject for ecological network science. In fact, macrophyte-animal networks offer an aquatic counterpart to terrestrial plant-animal networks. In this review, we show how the application of network analysis to aquatic macrophyte-animal associations has the potential to broaden our understanding of how coastal ecosystems function. Network analysis can also provide a key to understanding how such ecosystems will respond to on-going and future threats from anthropogenic disturbance and environmental change. For this, we: (i) identify key issues that have limited the application of network theory and modelling to aquatic animal-macrophyte associations; (ii) illustrate through examples based on empirical data how network analysis can offer new insights on the complexity and functioning of coastal ecosystems; and (iii) provide suggestions for how to design future studies and establish this new research line into network ecology., (© 2022 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Global change is striking harder and faster in the Mediterranean Sea than elsewhere, where high levels of human pressure and proneness to climate change interact in modifying the structure and disrupting regulative mechanisms of marine ecosystems. Rocky reefs are particularly exposed to such environmental changes with ongoing trends of degradation being impressive. Due to the variety of habitat types and associated marine biodiversity, rocky reefs are critical for the functioning of marine ecosystems, and their decline could profoundly affect the provision of essential goods and services which human populations in coastal areas rely upon. Here, we provide an up-to-date overview of the status of rocky reefs, trends in human-driven changes undermining their integrity, and current and upcoming management and conservation strategies, attempting a projection on what could be the future of this essential component of Mediterranean marine ecosystems., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

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., (© 2019 The Authors. Ecological Applications published by Wiley Periodicals, Inc. on behalf of Ecological Society of America.)

Marine protected areas (MPAs) represent the main tool for halting the loss of marine biodiversity. However, there is increasing evidence concerning their limited capacity to reduce or eliminate some threats even within their own boundaries. Here, we analysed a Europe-wide dataset comprising 31,579 threats recorded in 1692 sites of the European Union's Natura 2000 conservation network. Focusing specifically on threats related to marine species and habitats, we found that fishing and outdoor activities were the most widespread threats reported within MPA boundaries, although some spatial heterogeneity in the distribution of threats was apparent. Our results clearly demonstrate the need to reconsider current management plans, standardise monitoring approaches and reporting, refine present threat assessments and improve knowledge of their spatial patterns within and outside MPAs in order to improve conservation capacity and outcomes., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Research on regime shifts has focused primarily on how changes in the intensity and duration of press disturbances precipitate natural systems into undesirable, alternative states. By contrast, the role of recurrent pulse perturbations, such as extreme climatic events, has been largely neglected, hindering our understanding of how historical processes regulate the onset of a regime shift. We performed field manipulations to evaluate whether combinations of extreme events of temperature and sediment deposition that differed in their degree of temporal clustering generated alternative states in rocky intertidal epilithic microphytobenthos (biofilms) on rocky shores. The likelihood of biofilms to shift from a vegetated to a bare state depended on the degree of temporal clustering of events, with biofilm biomass showing both states under a regime of non-clustered (60 d apart) perturbations while collapsing in the clustered (15 d apart) scenario. Our results indicate that time since the last perturbation can be an important predictor of collapse in systems exhibiting alternative states and that consideration of historical effects in studies of regime shifts may largely improve our understanding of ecosystem dynamics under climate change., (© 2018 by the Ecological Society of America.)

Understanding how increasing human domination of the biosphere affects life on earth is a critical research challenge. This task is facilitated by the increasing availability of open-source data repositories, which allow ecologists to address scientific questions at unprecedented spatial and temporal scales. Large datasets are mostly observational, so they may have limited ability to uncover causal relations among variables. Experiments are better suited at attributing causation, but they are often limited in scope. We propose hybrid datasets, resulting from the integration of observational with experimental data, as an approach to leverage the scope and ability to attribute causality in ecological studies. We show how the analysis of hybrid datasets with emerging techniques in time series analysis (Convergent Cross-mapping) and macroecology (Joint Species Distribution Models) can generate novel insights into causal effects of abiotic and biotic processes that would be difficult to achieve otherwise. We illustrate these principles with two case studies in marine ecosystems and discuss the potential to generalize across environments, species and ecological processes. If used wisely, the analysis of hybrid datasets may become the standard approach for research goals that seek causal explanations for large-scale ecological phenomena., (© 2018 by the Ecological Society of America.)

Habitat-forming species sustain biodiversity and ecosystem functioning in harsh environments through the amelioration of physical stress. Nonetheless, their role in shaping patterns of species distribution under future climate scenarios is generally overlooked. Focusing on coastal systems, we assess how habitat-forming species can influence the ability of stress-sensitive species to exhibit plastic responses, adapt to novel environmental conditions, or track suitable climates. Here, we argue that habitat-former populations could be managed as a nature-based solution against climate-driven loss of biodiversity. Drawing from different ecological and biological disciplines, we identify a series of actions to sustain the resilience of marine habitat-forming species to climate change, as well as their effectiveness and reliability in rescuing stress-sensitive species from increasingly adverse environmental conditions., Competing Interests: The authors have declared that no competing interests exist.

Loss of algal canopies can result in a shift towards a turf-dominated state, where variability in species life-history traits can determine new mechanisms of feedback, and influence the degraded system under variable regimes of disturbance. By focusing on rockpools dominated by Cystoseira brachycarpa, we tested the hypothesis that the alga Dictyopteris polypodioides could take advantage of extreme regimes of disturbance related to storms, and outcompete other turfs through a distinctive combination of life traits. Replacement of the canopy was initially driven by a mix of taxon-specific life-traits and resulting assemblages were susceptible to intense events of disturbance. Subsequently, D. polypodioides dominated removal quadrats, favored by density-dependent abilities to intercept more light and reach larger size than the rest of turf. These new positive feedbacks may contribute to maintain the modified state of the system and influence its ability to withstand extreme abiotic conditions., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Developing early warning signals to predict regime shifts in ecosystems is a central issue in current ecological research. While there are many studies addressing temporal early warning indicators, research into spatial indicators is far behind, with field experiments even more rare. Here, we tested the performance of spatial early warning signals in an intertidal macroalgal system, where removal of algal canopies pushed the system toward a tipping point (corresponding to approximately 75% of canopy loss), marking the transition between a canopy- to a turf-dominated state. We performed a two-year experiment where spatial early warning indicators were assessed in transects where the canopy was differentially removed (from 0 to 100%). Unlike Moran correlation coefficient at lag-1, spatial variance, skewness, and spatial spectra at low frequency increased along the gradient of canopy degradation and dropped, or did not show any further increase beyond the transition point from a canopy- to a turf-dominated state (100% canopy removal). Our study provides direct evidence of the suitability of spatial early warning signals to anticipate regime shifts in natural ecosystems, emphasizing the importance of field experiments as a powerful tool to establish causal relationships between environmental stressors and early warning indicators., (© 2018 by the Ecological Society of America.)

Understanding how historical processes modulate the response of ecosystems to perturbations is becoming increasingly important. In contrast to the growing interest in projecting biodiversity and ecosystem functioning under future climate scenarios, how legacy effects originating from historical conditions drive change in ecosystems remains largely unexplored. Using experiments in combination with stochastic antecedent modelling, we evaluated how extreme warming, sediment deposition and grazing events modulated the ecological memory of rocky intertidal epilithic microphytobenthos (EMPB). We found memory effects in the non-clustered scenario of disturbance (60 days apart), where EMPB biomass fluctuated in time, but not under clustered disturbances (15 days apart), where EMPB biomass was consistently low. A massive grazing event impacted on EMPB biomass in a second run of the experiment, also muting ecological memory. Our results provide empirical support to the theoretical expectation that stochastic fluctuations promote ecological memory, but also show that contingencies may lead to memory loss., (© 2017 John Wiley & Sons Ltd.)

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.

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., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Gastropod assemblages from nearshore rocky habitats were studied over large spatial scales to (1) describe broad-scale patterns in assemblage composition, including patterns by feeding modes, (2) identify latitudinal pattern of biodiversity, i.e., richness and abundance of gastropods and/or regional hotspots, and (3) identify potential environmental and anthropogenic drivers of these assemblages. Gastropods were sampled from 45 sites distributed within 12 Large Marine Ecosystem regions (LME) following the NaGISA (Natural Geography in Shore Areas) standard protocol (www.nagisa.coml.org). A total of 393 gastropod taxa from 87 families were collected. Eight of these families (9.2%) appeared in four or more different LMEs. Among these, the Littorinidae was the most widely distributed (8 LMEs) followed by the Trochidae and the Columbellidae (6 LMEs). In all regions, assemblages were dominated by few species, the most diverse and abundant of which were herbivores. No latitudinal gradients were evident in relation to species richness or densities among sampling sites. Highest diversity was found in the Mediterranean and in the Gulf of Alaska, while highest densities were found at different latitudes and represented by few species within one genus (e.g. Afrolittorina in the Agulhas Current, Littorina in the Scotian Shelf, and Lacuna in the Gulf of Alaska). No significant correlation was found between species composition and environmental variables (r≤0.355, p>0.05). Contributing variables to this low correlation included invasive species, inorganic pollution, SST anomalies, and chlorophyll-a anomalies. Despite data limitations in this study which restrict conclusions in a global context, this work represents the first effort to sample gastropod biodiversity on rocky shores using a standardized protocol across a wide scale. Our results will generate more work to build global databases allowing for large-scale diversity comparisons of 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.

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.

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.

This study examined echinoderm assemblages from nearshore rocky habitats for large-scale distribution patterns with specific emphasis on identifying latitudinal trends and large regional hotspots. Echinoderms were sampled from 76 globally-distributed sites within 12 ecoregions, following the standardized sampling protocol of the Census of Marine Life NaGISA project (www.nagisa.coml.org). Sample-based species richness was overall low (<1-5 species per site), with a total of 32 asteroid, 18 echinoid, 21 ophiuroid, and 15 holothuroid species. Abundance and species richness in intertidal assemblages sampled with visual methods (organisms >2 cm in 1 m(2) quadrats) was highest in the Caribbean ecoregions and echinoids dominated these assemblages with an average of 5 ind m(-2). In contrast, intertidal echinoderm assemblages collected from clearings of 0.0625 m(2) quadrats had the highest abundance and richness in the Northeast Pacific ecoregions where asteroids and holothurians dominated with an average of 14 ind 0.0625 m(-2). Distinct latitudinal trends existed for abundance and richness in intertidal assemblages with declines from peaks at high northern latitudes. No latitudinal trends were found for subtidal echinoderm assemblages with either sampling technique. Latitudinal gradients appear to be superseded by regional diversity hotspots. In these hotspots echinoderm assemblages may be driven by local and regional processes, such as overall productivity and evolutionary history. We also tested a set of 14 environmental variables (six natural and eight anthropogenic) as potential drivers of echinoderm assemblages by ecoregions. The natural variables of salinity, sea-surface temperature, chlorophyll a, and primary productivity were strongly correlated with echinoderm assemblages; the anthropogenic variables of inorganic pollution and nutrient contamination also contributed to correlations. Our results indicate that nearshore echinoderm assemblages appear to be shaped by a network of environmental and ecological processes, and by the differing responses of various echinoderm taxa, making generalizations about the patterns of nearshore rocky habitat echinoderm assemblages difficult.

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.

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 use of hard coastal-defence structures, like breakwaters and seawalls, is rapidly increasing to prevent coastal erosion. We compared low-shore assemblages between wave-protected and wave-exposed habitats on breakwaters along a sandy shore of Tuscany (North-Western Mediterranean). Assemblages were generally characterized by a low diversity of taxa, with space monopolized by Mytilus galloprovincialis and Corallina elongata on the seaward side of breakwaters and by filamentous algae on the landward side. Assemblages in wave-protected habitats were characterized by greater temporal stability than those in exposed habitats and supported non-indigenous macroalgae such as Caulerpa racemosa and Codium fragile ssp. tomentosoides. Hence, the introduction of hard coastal-defence structures in otherwise soft-bottom dominated areas, attracting native and exotic rocky-bottom species, should be of great concern for the conservation of marine biodiversity at local and regional scales and for the management of biological invasions.

Neutral models and differential responses of species to environmental heterogeneity offer complementary explanations of species abundance distribution and dynamics. Under what circumstances one model prevails over the other is still a matter of debate. We show that the decay of similarity over time in rocky seashore assemblages of algae and invertebrates sampled over a period of 16 years was consistent with the predictions of a stochastic model of ecological drift at time scales larger than 2 years, but not at time scales between 3 and 24 months when similarity was quantified with an index that reflected changes in abundance of rare species. A field experiment was performed to examine whether assemblages responded neutrally or non-neutrally to changes in temporal variance of disturbance. The experimental results did not reject neutrality, but identified a positive effect of intermediate levels of environmental heterogeneity on the abundance of rare species. This effect translated into a marked decrease in the characteristic time scale of species turnover, highlighting the role of rare species in driving assemblage dynamics in fluctuating environments.

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.

Marine protected areas (MPAs) are increasingly used as a management tool to preserve species and habitats. Testing hypotheses about the effectiveness of MPAs is important for their implementation and to identify informative criteria to support management decisions. This study tested the general proposition that MPAs affected assemblages of algae and invertebrates between 0.0 and 0.5 m above the mean low water level of rocky coasts on two islands in the Tuscan Archipelago (northwest Mediterranean). Protection was concentrated mainly on the west coasts of the islands, raising the possibility that neither the full range of assemblages nor the relevant scales of variation were properly represented within MPAs. This motivated the comparison of assemblages on opposite sides of islands (habitats). The effects of MPAs and habitat were assessed with a multifactorial sampling design; hypotheses were tested about differences in structure of assemblages, in mean abundance of common taxa and in univariate and multivariate measures of spatial variation. The design consisted of three replicate shores for each condition of protected and reference areas on the west side of each island and three unprotected shores on the eastern side. Assemblages were sampled independently four times on each island between June 1999 and January 2001. At each time of sampling two sites were selected randomly at each of two tidal heights to represent midshore and lowshore assemblages on each shore. Estimates of abundance were obtained using non-destructive sampling methods from five replicate 20x20 cm quadrats at each site. Results indicated differences among habitats in structure of assemblages, in mean abundance of common taxa and in univariate and multivariate measures of spatial variation at the scale of shores. Most of these patterns were inconsistent with the predicted effect of management through MPAs. The data suggest that designation of MPAs in the Tuscan Archipelago should proceed through management of multiple shores and types of habitat selected to guarantee protection to a representative sample of assemblages and to the processes responsible for maintenance of spatial patchiness at different scales. This study also shows that considerations of spatial heterogeneity are important to underpin management decisions about the number, size and location of MPAs.