European Marine Protected Areas are well-placed and climate-resilient, but weakly protected

European Marine Protected Areas are well-placed and climate-resilient, but weakly protected | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article European Marine Protected Areas are well-placed and climate-resilient, but weakly protected Fabrice Stephenson, Patrick Eskuche-Keith, Anna Addamo, Peter Almond, and 11 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8894972/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted You are reading this latest preprint version Abstract Marine Protected Area (MPA) effectiveness depends on both coverage of species distributions and protection levels. It remains unclear whether Europe’s expanding MPA network adequately represents biodiversity now and as species redistribute under climate change. We provide the first pan-European assessment integrating MPA Guide protection levels with modelled present and future distributions for over 9,000 marine species. MPAs currently cover ~11 % of pan-European seas but encompass ~25% of species’ ranges, indicating preferential placement in biodiversity-rich areas. Representation remains stable under end-century projections (SSP2-4.5), and species at high climate risk generally retain or increase MPA coverage, suggesting protection of climatic refugia. However, < 1 % of species’ distributions fall within areas restricting harmful practices such as fishing. Europe’s MPA network is well placed for biodiversity conservation and climate resilience but weakly protected, limiting its conservation impact and highlighting the need to strengthen management and strategically expand effectively protected and climate-resilient areas. Biological sciences/Ecology/Conservation biology Biological sciences/Ecology/Biogeography Biological sciences/Ecology/Biodiversity Earth and environmental sciences/Ocean sciences/Marine biology Scientific community and society/Scientific community/Policy Species distribution modelling marine spatial planning area-based conservation range shifts management effectiveness marine policy Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Globally, marine biodiversity is declining at an unprecedented rate, driven primarily by human activities that further degrade habitats, pollute ecosystems, and intensify resource extraction 1,2 . Climate change adds to these pressures by raising ocean temperature and consequently changing species distributions, with thousands of marine species having already shifted their distribution poleward 3,4 . Together, these drivers transform marine ecosystems at rates frequently exceeding the practical capacity of existing management systems. Marine Protected Areas (MPAs) are effective tools for reducing cumulative stressors, safeguarding biodiversity, and sustaining ecosystem services 5-7 . By protecting locations likely to remain suitable under future climates, MPAs can support ecological resilience 8 . These attributes underpin global conservation commitments, including targets of the Global Biodiversity Framework and the recent High Seas Treaty 9 . However, expansion of MPA networks alone does not guarantee biodiversity protection or climate readiness; effectiveness depends on placement, restricted pressures, enforcement, and stakeholder participation 10 . Despite growing global MPA coverage, evidence suggests that current networks insufficiently represent and protect marine biodiversity. Only a minority of species have substantial range fractions within existing MPAs 11-13 , and most designated sites continue to allow ecologically damaging activities, such as industrial fishing including seabed trawling 14,15 , eroding intended conservation outcomes 16,17 . To address inconsistent definitions and expectations of protection, the MPA Guide provides a framework linking protection levels to expected ecological outcomes 6 . Its global application revealed that only one-third of the 100 largest MPAs (90 % of global MPA coverage, representing 7.3 % of global ocean area) qualify as ‘Fully’ or ‘Highly’ protected 18 , underscoring the disparity between nominal protection and realised ecological security (the so-called ‘paper parks’ phenomenon). Across Europe, conservation ambition is shaped both by global commitments and also by a complex patchwork of regional and national governance frameworks. European Union (EU) legislation, notably the Birds and Habitats Directives 19,20 , supports the Natura 2000 network, while regional conventions (e.g., OSPAR, the Helsinki Convention, the Bucharest Convention and the Barcelona Convention) complement EU policy across basin scales 21-24 . However, these frameworks historically emphasise spatial designation rather than ecological representativity or specific management requirements, producing a heterogeneous mosaic of MPA types and regulatory regimes. This complexity complicates assessments of true protection levels and coordination of coherent, climate-ready conservation across jurisdictions. Within EU waters, MPAs cover >12 % of marine areas, yet only 1.5 % is classified as ‘strongly protected’ (i.e., falling within the ‘Fully’ or ‘Highly’ protected categories under the MPA Guide) 6,25 . The EU’s Biodiversity Strategy for 2030 aims to designate 30 % of Earth’s ocean area as protected areas by 2030 (“30 × 30”) and ensure at least 10 % of European seas are ‘strictly’ protected (i.e., left essentially undisturbed from human pressures and threats, corresponding to full protection as defined in the MPA guide) 6,26,27 . The EU’s ambition will require both substantial expansion and the effective management of MPAs capable of delivering strong ecological outcomes 28,29 . Critically, assessments of protection levels have not extended to non-EU European marine nations, including Iceland, Norway, the Faroe Islands, and the United Kingdom, which collectively manage a large share of pan-European seas and are ecologically connected. Consequently, the actual extent of marine protection in Europe remains unclear. A central uncertainty is whether existing MPA locations adequately capture marine species’ distributions, both under present-day conditions and as ranges shift under climate change. As species ranges expand or contract due to shifting conditions 30 , increasing proportions of their ranges may fall outside or inside current MPA boundaries, raising the question of the adequacy of existing networks 31-33 . Recent advances in open-access biodiversity and environmental datasets, such as OBIS 34 and Bio-ORACLE v3 35 , together with high-resolution species distribution models (SDMs) for thousands of marine taxa 36 , enable forecasts of future species distributions under alternative emissions scenarios 37 . These developments offer opportunities to evaluate species coverage by MPAs under climate change. Here we assessed how MPAs with protection levels classified using the MPA Guide overlapped with predicted distributions for 9,794 marine species across pan-European seas; defined as the Exclusive Economic Zones of EU member states, Iceland, Norway (including Svalbard and Jan Mayen), the Faroe Islands, and the United Kingdom. We quantified the proportion of current species distributions encompassed by MPAs across protection levels and major biotic groups, and evaluated climate readiness by comparing present-day distributions with those projected for the end of the century (2090-2100) under a moderate ‘most likely’ Shared Socio-economic Pathway emissions scenario (SSP2-4.5) 38 . We categorised species into climate-risk classes based solely on projected changes in range size (e.g. 32 ), whereby ‘High climate-risk’ species were projected to lose > 50 % of their present-day distribution, ‘Medium climate-risk’ species were projected to lose 10-50 % of their distribution, and ‘Low climate-risk’ species were projected to lose <10 % or even gain distribution area. We examined whether MPAs were positioned to protect these classes as climatic conditions change. Through this analysis, we provide the first pan-European assessment of the effectiveness and climate readiness of current marine protection across all European jurisdictions. Results Distribution of MPAs by protection level MPAs covered 1,017,565 km², approximately 11 % of pan-European seas. Most (91 %) offered weak protection (i.e., ‘Light’, ’Minimal’ or ‘Incompatible’), while only ~1 % were strongly protected (i.e., ‘Fully’ or ‘Highly’), with the remainder unclassified (Figure 1, Table 1). Individual national Exclusive Economic Zones (EEZs) showed considerable variability, with MPA coverage ranging from <1 % to ~48 %, although all countries had <1 % of their EEZ under strong protection (Table S1). Coverage of species ranges by MPA protection level Across all species, median MPA coverage of distributions was 25 % (Figure 2, Table S2). Among the 4,917 species with ≥ 25 % of their ranges falling within MPA boundaries (i.e., ≥ overall median), median coverage by strongly protected MPAs was < 1 % (95th percentile = 5 %). Overall, across all species, strongly protected MPAs generally covered < 1 % of species’ distributions (median <1%), whereas weakly protected MPAs accounted for most coverage (median = 22 %; Table S2). Across biotic groups, median coverage of species distributions by all MPAs ranged from 14 % (pelagic invertebrates) to 30 % (demersal fish) (Figure 2, Table S2). Median coverage by MPAs with strong protection levels remained < 1 % for all groups, with 95th percentiles < 2 % in all cases (Table S2). Coverage by weak protection levels ranged from 12 % (pelagic fish and invertebrates) to 25 % (benthic invertebrates). Median coverage of threatened and habitat-forming species distributions across all MPAs was 29 % and 25 %, respectively (Table S3). Among threatened species (i.e., IUCN Red List Vulnerable, Endangered or Critically endangered species), two thirds ( n = 146) had ≥ 25 % of their individual species’ distribution protected. For habitat-formers (biogenic reefs, canopy-forming macroalgae and seagrasses), half of the species ( n = 62) met the threshold of ≥ 25 % protection. Median coverage by strongly protected MPAs was < 1 % for both threatened and habitat-forming species, while weakly protected MPAs accounted for substantially higher coverage (26 % and 22 %, respectively) (Figure 3, Table S3). Table 1: Total areas (km 2 ) covered by each MPA protection level. Values in brackets express this as a percentage of the total area covered by MPAs in each column. Combined areas are presented for the Icelandic, Faroese, Norwegian and UK assessments and also for the full pan-European study region. For the Icelandic and Faroese MPAs, a lack of sufficient information for assessment meant these were recorded as Unclassified. The area of each protection level as a percentage of the total area of pan-European seas is also presented. Combined protection level MPA Guide protection level UK, Norway, Iceland and Faroes MPAs pan-European study region Percentage of pan-European seas (%) Strong Fully 57 (0.1) 1,156 (0.1) < 0.1 Highly 1,003 (0.2) 8,339 (0.8) 0.1 Weak Lightly 24,507 (6.0) 110,737 (10.0) 1.2 Minimally 170,850 (41.5) 473,584 (46.5) 5.2 Incompatible 221,746 (51.5) 342,543 (33.7) 3.7 Unclassified 3,361 (0.8) 81,207 (8.0) 0.9 TOTAL 411,525 1,017,565 11.1 Impacts of climate change on species distributions A marginally smaller subset of spatial predictions ( n = 9,635) was available under both present-day environmental conditions and the SSP2-4.5 scenario, due to filtering of SDM outputs during the data processing stages (see Methods). These were used to investigate potential future range changes. Approximately 54 % of these species ( n = 5,227) were projected to experience net decreases in their distribution area within pan-European waters, while ~45 % ( n = 4,357) were projected to experience net increases and 51 species were not projected to show a net area change. There was high variability in projected changes in distribution area across biotic groups. Primary producers and both pelagic and benthic invertebrates were dominated by species projected to experience decreases in distribution area within the study region, with median losses of 7 %, 6 % and 3 %, respectively. In contrast, the majority of elasmobranchs were projected to experience distribution area increases, with median gains of 7 %, while mammals and demersal and pelagic fish were projected to experience median increases of ~ 2 % overall. ‘High climate-risk’ species (i.e., defined in this study as those projected to lose > 50 % of their present-day distribution area under climate change) comprised ~18 % of taxa ( n = 1,723), while ‘Medium climate-risk’ species (10-50 % loss) accounted for ~25 % ( n = 2,404). The remaining ~57 % of taxa ( n = 5,508) were classed as at ‘Low climate-risk’ (< 10 % loss or net gains). Impacts of climate change on coverage of species distributions by MPA protection levels When all species were aggregated, the median projected changes in range coverage from MPA protection levels by 2100 under SSP2-4.5 were negligible (< 1 % in either direction) (Table S4). Similarly, when split by biotic group, median change across all MPAs ranged from -1 % (elasmobranchs) to +1 % (mammals) (Table S4). However, changes in coverage varied widely across risk classes and among groups. For ‘High climate-risk’ species, median relative change in MPA coverage under SSP2-4.5 was positive across all biotic groups, with increases between < 1% (demersal fish and invertebrates) and ~12 % (mammals) (Figure 4, Table S5). Median changes in the proportion of distributions covered by strongly protected MPAs were minor (< 1 % gains) for all groups while median changes in coverage within weakly protected MPAs ranged from < -1% (demersal fish) to +11 % (mammals) (Table S5). For ‘Medium climate-risk’ taxa, the majority of biotic groups were projected to gain proportional coverage by MPAs, up to ~3 % (mammals), although benthic invertebrates were generally projected to lose coverage (median: -1%) (Figure S1, Table S6). Median changes in coverage by strongly protected MPAs were minor (< 1 % in either direction), while median changes in coverage by weakly protected MPAs ranged from < -1 % (benthic invertebrates) to +3 % (mammals) (Table S6). For ‘Low climate-risk’ taxa, most species in all groups were projected to experience limited decreases in the proportion of their distributions covered by MPAs (Figure S2, Table S7). Median changes were generally negative but minor, with coverage changes by both strongly and weakly protected MPAs ranging from -1 % to no change (Table S7). Discussion We provide the first pan-European evaluation of MPA effectiveness integrating predicted distributions for 9,794 marine species with up-to-date MPA protection level classifications. Despite occupying only ~11 % of pan-European seas, we show that MPAs generally encompass one-quarter of species’ distributions, suggesting they are broadly well placed to protect biodiversity. Encouragingly, these conservation benefits persist under future climate conditions; median coverage changes are negligible (< 1 % across taxa), suggesting the current spatial configuration of MPAs is robust to projected range shifts under an intermediate emissions scenario (SSP2-4.5). However, strong protection remains extremely limited, with < 1 % of species’ distributions falling within Fully or Highly protected areas. Together, these findings highlight the potential of Europe’s MPA distribution and the urgent need to strengthen protection levels to secure biodiversity outcomes now and in the future. MPAs are well-placed, but not optimised The large proportion of species’ ranges within MPAs relative to their areal footprint indicates their preferential placement in biodiversity-rich regions. This may reflect deliberate prioritisation of ecologically important areas or the concentration of many early designations in high-richness coastal zones 39 . Such patterns are consistent with the idea that spatial planning efforts, whether explicit or emergent, have captured areas of relatively high conservation value 40,41 . Because existing MPAs already overlap substantial parts of many species’ distributions, strengthening protection and management within current boundaries, particularly in sites with high biodiversity representation, could deliver disproportionately strong conservation outcomes without relying solely on further spatial expansion of MPAs 6,42 . Systematic conservation planning has repeatedly shown that alternative configurations can achieve similar or higher representation at lower area cost, including in global analyses using tools such as Zonation 13,43,44 . Our results highlight that while pan-European MPAs are generally well placed (although weakly protected), there remains scope to refine planning for more even representation across taxa and marine regions. For example, pelagic invertebrates and fishes have lower median MPA coverage (14 % and 15 %, respectively) than other groups and substantially lower than the overall median (25 %). This likely reflects these species often having larger ranges than their benthic and demersal counterparts 45,46 . In some cases, this underrepresentation may be less problematic; zooplankton, for instance, are not typically subject to direct fishing pressure, so lower MPA coverage may not equate to major gaps in protection 47 . This may also apply to highly migratory species including certain elasmobranchs or marine mammals, where protecting strategic places (e.g. spawning, nursery and feeding grounds) provides greater benefits than simply increasing overall overlap with MPAs 48,49 . By contrast, for groups exposed to intense anthropogenic impacts, such as many widespread pelagic fishes or benthic invertebrates sensitive to bottom-contact fishing, low representation within MPAs could indicate missed opportunities for targeted improvements in spatial coverage and management 50 . Climate-driven change and the resilience of spatial placement Projected changes in species distributions reveal some climate-driven redistribution across pan-European seas by the end of the century. Under SSP2-4.5, approximately 54 % of species are expected to shrink their geographic range within European waters, with ~18 % classified here as ‘High climate-risk’ due to declines in distribution area exceeding 50 %. These projections are specific to the pan-European seas, yet in many cases likely reflect broader poleward or basin-scale redistributions under warming 51-54 . Range changes are uneven across functional groups, with invertebrates and primary producers projected to display median relative losses, whereas marine mammals, fish and elasmobranchs are projected to experience relative gains. Despite projected range shifts, the proportion of species’ distributions falling inside MPAs remains remarkably stable in future. Across all biotic groups and protection levels, median changes in MPA coverage are < 1 %, indicating that current MPAs continue to intersect substantial portions of species’ distributions, at least under an intermediate emissions scenario. ‘High climate-risk’ species show a median increase in their representation within MPAs despite losing large fractions of their total pan-European range, implying that most projected loss occurs outside protected areas while currently suitable habitat within MPAs persists. Conversely, ‘Low climate-risk’ species, particularly those projected to expand their distributions, tend to experience declining representation, reflecting that most MPAs were not designed with future range expansions or poleward shifts in mind 10 . Together, these patterns suggest pan-European MPA placement exhibits substantial climate robustness, with protected areas acting as refugia for species undergoing regional declines, consistent with observations from other marine systems 55 . Our measure of ‘climate-risk’ provides a simple indicator which could be used to prioritise certain taxa based on their expected range changes. In addition to proportional area loss however, more detailed case studies and assessments should account for overall range size as this is a major predictor of extinction risk under climate change 56 . These findings resonate with broader work linking spatial planning to climate-resilient conservation. Designing MPAs solely around present-day habitat suitability is not guaranteed to safeguard species under climate change, whereas integrating projected distributions may improve protection of key climate refugia 32 . In some large MPAs, species representation is projected to increase under climate change, underscoring that existing protections can remain relevant as conditions shift 39 . However, threatened and commercially important species in pan-European MPAs face changing climate conditions, particularly under high-emission scenarios and in semi-enclosed seas such as the Mediterranean, Baltic and Black seas 50,57 . In these regions, even dense MPA networks may struggle to offset climate-driven changes in environmental suitability, unless protection levels are sufficient to safeguard biodiversity 42 . Protection quality is the critical bottleneck The most pervasive weakness of present European MPAs lies not in their location but in how strongly they are protected. Across all taxa, fewer than 1 % of species’ distributions occur within strongly protected MPAs. This means that even species with extensive spatial overlap with MPAs are subject to weak restrictions on extractive activities and other pressures. These results support the conclusion that management shortfalls and low protection levels, rather than spatial gaps, are the primary barriers to delivering the biodiversity outcomes promised by MPA policy 5,6,16,25,58 . The poor representation of strongly protected MPAs within all individual EEZs highlights that this is a widespread issue which all relevant European nations need to take legislative action on in order to truly safeguard their marine biodiversity. At the regional scale, improving MPA protection levels would also provide a strategic contribution towards achieving the European 10 % target for strict protection (corresponding to full protection as defined in the MPA guide) 6,59,60 . Our analysis considered protection levels but not the stage of establishment, which can be a critical determinant of ecological performance 6 . Implementation and active management are key components of the MPA Guide and can be critical for MPA effectiveness 6,61 . Actively managed MPAs deliver significantly stronger ecological benefits than sites that remain partially implemented or, worse, are only designated on paper 62 . Thus, increasing the area considered as Fully and/or Highly protected will be necessary but not sufficient without full implementation, enforcement, and adequate resources for management. However, the overwhelming majority of MPAs within pan-European seas have low designated levels of protection, which, even if implemented or actively managed, may provide no measurable ecological benefits and could even have negative impacts (as measured using indicators such as fish density or biomass) 62 . It is therefore imperative that efforts to improve the effectiveness of existing pan-European MPAs focus on both their active management and the prohibition of harmful or destructive activities within their boundaries. The interaction between climate risk and protection levels is particularly relevant. ‘High climate-risk’ species are projected to maintain or increase the proportion of their range within MPAs under future conditions, but their low coverage by strong protection undermines the potential benefits of this favourable spatial configuration 50 . Variation among taxa further emphasises the need to align protection levels with ecological need. Groups with limited exposure to direct anthropogenic impacts, such as pelagic invertebrates, may not require extensive strong protection 63 . By contrast, benthic invertebrates, elasmobranchs, and commercially targeted fishes face intense fishing pressure yet receive disproportionately weak protection 25,50 . For these taxa, the scarcity of fully or highly protected areas directly compromises prospects for recovery, resilience, and long-term viability 51 . Taken together, our findings indicate that simply increasing the areal extent of MPAs will not address the core challenges facing marine biodiversity. Expansion alone cannot compensate for inadequate protection levels 16,29,62 or limited management enforcement 6,42 . Limitations and future directions Our analysis is subject to several limitations that should inform interpretation and future work. First, we relied on species distribution models derived from available biodiversity and environmental data; these models inevitably carry uncertainty related to limited and uneven sampling across species’ geographic ranges, choices of model algorithms and parameters, and accuracy of climate projections 36,37 . Second, we focused on a single intermediate emissions scenario (SSP2-4.5) representing a ‘most-likely’ future under current global socio-economic trajectories. Higher-emission pathways are likely to produce greater range extensions and contractions in response to changing climate, particularly in enclosed and high-latitude seas 54 . Addressing these points can be accommodated by monitoring actual changes in species distribution and abundance and adapting conservation accordingly. Our analyses were restricted to EEZs and did not consider areas beyond national jurisdiction (ABNJ), which are increasingly important in European and global conservation strategies. With the recent implementation of the High Seas Treaty, there is a clear basis for expanding similar analyses to ABNJ to support systematic spatial prioritisation and conservation planning there. Conclusion Our results indicate that strengthening protection within the existing MPA network (alongside carefully planned expansion) offers one of the most direct and effective pathways for Europe to meet its marine biodiversity commitments. Major conservation benefits could be realised by upgrading the protection level and ensuring active management of current sites, particularly those that already capture a large share of species’ distributions or act as climate refugia, while using new designations to fill residual spatial and taxonomic gaps. In this context, the drive to achieve “30 × 30” should not only increase the areal extent of MPAs, but must prioritise Fully and Highly protected areas and ensure that all MPAs are effectively managed, ecologically coherent and socially equitable (e.g. 28,29 ). Furthermore, these MPAs are likely to help restore marine ecosystems and the recovery and long-term sustainability of fisheries 64,65 . Our pan-European assessment adds to growing evidence that the quality and level of protection within MPAs will be critical in determining whether quantitative targets such as protecting 30 % of European seas translate into meaningful, durable outcomes for both biodiversity, blue economies and people in a changing ocean. Methods Study region and MPAs This study investigated protection levels at the pan-European scale, considering the EEZs of EU member states in addition to those of the Iceland, Norway (including Svalbard and Jan Mayen under Norwegian sovereignty), the Faroe Islands, and the United Kingdom (Figure 1). The resulting study region covered a total area of approximately 9,160,000 km 2 . Polygon shapefiles representing MPA boundaries were obtained for each constituent country: for EU member states from 25 , originally extracted from the European Environment Agency 66 ( n = 5,362); for the UK, polygons representing marine-associated Special Areas of Conservation (SAC), marine-associated Special Protection Areas (SPA), Nature Conservation Marine Protected Areas (NCMPA), and Marine Conservation Zones (MCZ) (including Highly Protected Marine Areas (HPMA)) ( n = 391) were obtained from various geospatial platforms of the UK's Statutory Nature Conservation Bodies (SNCBs) 67 ; Protected area shapefiles for Norway ( n = 858) were obtained from 68 , while shapefiles for Iceland ( n = 68) and the Faroe Islands ( n = 3) were obtained from the World Database on Protected Areas (WDPA) 69 and clipped to exclude areas on land. MPA protection level assignments The MPA Guide decision tree framework 6,59 was applied to classify each MPA or zone into one of four protection levels or as incompatible with conservation, based on regulations for seven activity types: mining, dredging and dumping, infrastructure, anchoring, fisheries, aquaculture and non-extractive uses 6 . The protection levels were: ‘Fully’ protected, where all extractive or destructive activities are prohibited and other impacts are minimized; ‘Highly’ protected, where only light extractive uses with low total impact are permitted; ‘Lightly’ protected, where some biodiversity protection exists, but moderate extractive impacts are allowed; ‘Minimally’ protected, allowing extensive extraction and high impact uses; Incompatible with conservation of nature, whereby activities deemed too impactful for biodiversity protection are allowed to occur. MPAs lacking sufficient regulatory information were labelled Unclassified . ‘Fully’ and ‘Highly’ protected areas are expected to deliver the strongest biodiversity and ecosystem recovery benefits, with ‘Fully’ protected MPAs offering the greatest gains, whereas ‘Lightly’ and ‘Minimally’ protected MPAs may yield limited or no species-specific benefits, with little effect on biodiversity at the ecosystem level, for example due to trophic cascades 59,62,70 . For EU member states, protection level classifications were already available from 25 .Aminian-Biquet et al., 2024 The EU MPA protection levels were available for two impact scenarios to account for situations where there was uncertainty regarding the scale and potential impacts of activities: the first assumed that, unless explicitly specified, the lowest possible impacts were occurring, while the second assumed the highest possible impacts were occurring 25 . For this study, assigned MPA protection levels assuming the lowest possible impacts were used, therefore in cases where there was uncertainty regarding the scale and potential impacts of activities the results represent an optimistic view of the levels of protection provided by the MPAs. Sufficient information was available for most UK and Norwegian MPAs to make informed decisions regarding the levels of impact for different activities, but where uncertainty did exist, the same approach was followed as for the EU MPAs 25,67,68 . There was insufficient information to assess the levels of protection provided by the Iceland and the Faroe Islands protected areas, therefore these were all categorised as ‘Unclassified’. Combined, the Icelandic and Faroese areas represent only 3,488 km 2 (~0.3% of the total MPA area considered in this study) but the total marine area represented by their constituent national EEZs are approximately 11% of pan-European seas, therefore it was important to consider them in the analyses. All overlapping MPA polygons (i.e., resulting from multiple designations) with the same protection level were merged to remove areas of overlap and thereby avoid the double-counting of areas. In cases where overlapping MPA polygons represented different protection levels, the highest level of protection present was assigned to the overlap. Species distribution modelling The current distributions of marine species within national EEZs were obtained from species distribution models (SDMs) in 36 . These SDMs were developed using three different algorithms (Maxent, Random Forest, and XGBoost) under a point-process framework, and are also available for different time periods and Shared Socio-economic Pathway emissions scenarios. For the present analysis, we extracted species distributions for the present-day and for the ‘most-likely’ SSP2-4.5 scenario for the year 2100. Where possible, the ensemble prediction was selected, but where the ensemble was not available the model with the highest performance across all methods (as measured by the Continuous Boyce Index, 71 ) was selected. Species distribution layers were then clipped and masked to the area where the species was considered native. This was done by using the biogeographic realms from 72 ; realms were overlaid with occurrence records, and those containing records were considered part of the native range. A buffer of 0.2 degrees was applied to the occurrence records to ensure that realms at the edges of species distributions were also included. We further restricted the distributions to the maximum recorded depth for each species, based on the occurrence data used to fit the models. The same process was applied to the future prediction layers. Predictions were also uncertainty-discounted by subtracting the standard deviation across bootstraps from the average predicted probability of occurrence for each cell, using a fixed weight (0.5). Finally, we applied a threshold to convert probability of occurrence to presence or absence, using the model-derived threshold that maximized sensitivity (i.e. correctly identifying presences) and specificity (i.e. correctly identifying absences). This threshold has been shown to perform well even with presence-only models, such as this study, where no absence data is available and only background points are used 73,74 . As a final screening step, species with fewer than 50 predicted raster cells under present-day conditions were excluded from further analysis to avoid including taxa that are marginal or poorly predicted within the study area. Appropriate present-day distribution predictions were ultimately available for 9,794 species. Projected distributions under SSP2-4.5 were also extracted for these taxa, although slightly fewer were available ( n = 9,635) due to the processing steps above. Biotic group assignments Species were classified into primary biotic groups based on their taxonomy and ecology. All photoautotrophs including macrophytes (e.g., seagrasses and macroalgae) and phytoplankton were assigned to the ‘Primary producers’ group. Demersal and pelagic habitat assignments for fish were obtained based on their classification in FishBase (‘Pelagic fish’ were those species identified as ‘benthopelagic’, ‘bathypelagic’, ‘pelagic-oceanic’, ‘pelagic-neritic’, or ‘pelagic’; ‘Demersal fish’ were those species identified as reef-associated’, ‘demersal’, or ‘bathydemersal’), using the ‘ rfishbase ’ r package 75 . Invertebrates were split into either benthic or pelagic based on their taxonomy and known ecology in a top-down manner from class to species level: at each level, the relevant functional group classification from WoRMS was assigned to all taxa within that level (if available); assignments for any remaining taxa were then conducted at the next level (e.g. order), and so on until family level. For the few remaining taxa, assignments were then made at the species level. The invertebrates were then also screened to identify parasitic species, which were assigned to an ‘Others’ group. The invertebrate order Limnomedusae (Class Hydrozoa) was also assigned to the group ‘Others’ due to their combined benthic and pelagic lifecycle. Taxa deemed not to be strictly marine (birds and marine-associated terrestrial invertebrates), and reptiles (all turtles) were also assigned to this group. Due to the mixed membership of this ‘Others’ group, all constituent species ( n = 1,028) were excluded from further analyses. To explore how well MPA classes cover the distributions of key taxa, species were also separately grouped based on their conservation status and whether they are important habitat-formers. Species were classified as either threatened or non-threatened, with the former made up of those listed on the IUCN Red List as Vulnerable, Endangered or Critically endangered (see further details in 76 ). The habitat forming species were those categorized into one of nine distinct biogenic habitat groups: bryozoan reefs; cold-water coral reefs; coralligenous platforms; macroalgae forests; deep-sea sponge grounds; mollusc reefs; polychaete reefs; seagrasses; and shallow-water sponge reefs 77 . Analyses Overlap between MPA protection levels and species All analyses were conducted in R version 4.5.0 78 . Rasters were handled using the terra package 79 . For each species, the total area of overlap between cells with predicted presences and each MPA protection level was determined using the exactextractr package 80 . This approach facilitates the calculation of zonal statistics for rasters covered by polygon areas, including areas of partial overlap 80 . The proportion of each species’ range (i.e., the summed area of cells with predicted presences) covered by each MPA protection level was then calculated. This process was conducted for the predicted spatial distributions under both present-day and future (SSP2-4.5) environmental conditions. The distributions of MPA protection level coverage across all species and for each biotic group (including threatened and habitat-forming taxa) under present-day environmental conditions were visualised using violin plots and boxplots, and described using summary statistics (medians, 95 th percentiles and minimum/maximum). When reporting results in text, the two highest protection levels (‘Fully’ and ‘Highly’) are referred to as a single combined level strong protection , and the remaining levels (excluding ‘Unclassified’) were combined into a weak protection level, following 25 . This was done to clearly differentiate the protection levels which are the most likely to have positive outcomes for biodiversity 6 . Protection levels are reported individually in figures except for ‘Fully’ and ‘Highly’ which represent low overall areas (~1% of all pan-European MPAs) and were therefore grouped into “Fully / Highly” for visual clarity. Summary statistics in tables are reported separately for each MPA protection level. To explore the climate readiness of the pan-European MPAs, the overlap of future species distributions with MPAs was calculated using the same approach as that described for present-day distributions. The relative changes in the areas of overlap for each species and MPA protection level between current predicted distributions and projected future distributions were also calculated, by dividing the future projected areas of coverage by the current day predicted areas of coverage. As a simple indication of how climate change may impact species both directly through expansions or contractions in total range area and indirectly through shifts in distribution and associated coverage by MPAs, species were split into risk categories based on their relative changes in total range area (e.g., see 32 ): species projected to experience distribution contractions of >50% were considered ‘High climate-risk’; species projected to lose between 10% and 50% of their distribution were considered ‘Medium climate-risk’; those projected to lose <10% of their distribution or experience net gains were considered ‘Low climate-risk’. The distribution of relative changes in areas of overlap with MPA protection levels was summarised by biotic group for each risk class separately. Declarations Data availability Data and code will be made available in an open-access repository (e.g. Zenodo) upon acceptance of this manuscript. Acknowledgements F.S. and P.A.E-K were funded through the Newcastle University One Planet NUAcT Fellowship. F.S., P.A., and D.S. acknowledge support from Oceana for work related to Marine Protected Areas in a Changing Climate. A.M.A., S.C.P., J.A., and M.J.C. acknowledge support from the MPA Europe (HORIZON-CL6-2023-BIODIV-01-12, Grant Agreement No. 101059988), co-funded by the European Union’s Horizon Europe Programme. B.H.C., I.S., J.M.A., E.M., and A.R.K. acknowledge support from the MARHAB project (HORIZON-CL6-2023-BIODIV-01-4, Grant Agreement No. 101135307), funded by the European Union’s Horizon Europe Programme. B.H.C., I.S., and J.A. also received support from Portuguese national funds from FCT – Foundation for Science and Technology through contracts UID/04326/2025 (DOI: https://doi.org/10.54499/UID/04326/2025), UID/PRR/04326/2025 (DOI: https://doi.org/10.54499/UID/PRR/04326/2025), and LA/P/0101/2020 (DOI: https://doi.org/10.54499/LA/P/0101/2020) to CCMAR. B.H.C. also acknowledges support from the MarPlus2024 (Algarve 2030, ALGARVE-FSE+-01177700) project. The authors also thank Beth Pike, Jenna Sullivan-Stack, Kirsten Grorud-Colvert, and Joachim Claudet for valuable discussions. CRediT statement Patrick A. Eskuche-Keith: Conceptualisation, Data Curation, Methodology, Investigation, Formal analysis, Visualisation, Writing - original draft, Writing - Review & Editing Fabrice Stephenson: Conceptualisation, Supervision, Funding acquisition, Methodology, Investigation, Writing - original draft, Writing - Review & Editing Mark J. Costello: Conceptualisation, Supervision, Funding acquisition, Methodology, Writing - original draft, Writing - Review & Editing Pippa J. Moore: Conceptualisation, Funding acquisition, Writing - Review & Editing Bárbara Horta e Costa: Conceptualisation, Funding acquisition, Supervision, Data Curation, Methodology, Writing - original draft, Writing - Review & Editing Anna M. Addamo: Conceptualisation, Methodology, Data Curation, Writing - original draft, Writing - Review & Editing Silas C. Principe: Conceptualisation, Methodology, Data Curation, Formal analysis, Writing - original draft, Investigation, Writing - Review & Editing Peter M. Almond: Writing - original draft, Writing - Review & Editing Jorge Assis: Conceptualisation, Writing - Review & Editing Inês Sousa: Investigation, Data Curation, Writing - Review & Editing Even Moland: Investigation, Data Curation, Writing - Review & Editing Alf R. Kleiven: Investigation, Data Curation, Writing - Review & Editing Johanna M. Aarflot: Investigation, Data Curation, Writing - Review & Editing Elsa Sim: Investigation, Data Curation, Writing - Review & Editing Daniel Skerritt: Conceptualisation, Writing - Review & Editing References Jaureguiberry, P. et al. The direct drivers of recent global anthropogenic biodiversity loss. Science Advances 8 (2022). https://doi.org/10.1126/sciadv.abm9982 Halpern, B. S., Frazier, M., O’Hara, C. C., Vargas-Fonseca, O. A. & Lombard, A. T. Cumulative impacts to global marine ecosystems projected to more than double by mid-century. Science 389 , 1216-1219 (2025). https://doi.org/10.1126/science.adv2906 Chaudhary, C., Richardson, A. J., Schoeman, D. S. & Costello, M. J. Global warming is causing a more pronounced dip in marine species richness around the equator. Proceedings of the National Academy of Sciences 118 , e2015094118 (2021). https://doi.org/10.1073/pnas.2015094118 He, Q. & Silliman, B. R. Climate Change, Human Impacts, and Coastal Ecosystems in the Anthropocene. Current Biology 29 , R1021-R1035 (2019). https://doi.org/10.1016/j.cub.2019.08.042 Costello, M. J. Long live Marine Reserves: A review of experiences and benefits. Biological Conservation 176 , 289-296 (2014). https://doi.org/10.1016/j.biocon.2014.04.023 Grorud-Colvert, K. et al. The MPA Guide: A framework to achieve global goals for the ocean. Science 373 (2021). https://doi.org/10.1126/science.abf0861 Gonçalves, E. J. Marine Protected Areas as Tools for Ocean Sustainability, 131-141, Springer International Publishing (2023). White, J. W. et al. Measurements, mechanisms, and management recommendations for how marine protected areas can provide climate resilience. Marine Policy 171 , 106419 (2025). https://doi.org/10.1016/j.marpol.2024.106419 Stephens, T. The Kunming–Montreal Global Biodiversity Framework. International Legal Materials 62 , 868-887 (2023). https://doi.org/10.1017/ilm.2023.16 Wenzel, L. et al. Establishing Marine Protected Areas in a Changing Climate. IUCN WCPA Technical Report Series No. 9., (2025). Klein, C. J. et al. Shortfalls in the global protected area network at representing marine biodiversity. Scientific Reports 2015 5:1 5 (2015). https://doi.org/10.1038/srep17539 Appeltans, W. et al. Biodiversity knowledge and threats on marine life: assessing no-take zones as a refuge for marine species. (IOC-UNESCO, 2024). Zhao, Q. & Costello, M. J. Ecologically representative Marine Protected Area planning can think globally and act locally. Trends in Ecology & Evolution 40 , 772-781 (2025). https://doi.org/10.1016/j.tree.2025.05.007 Edgar, G. J. et al. Global conservation outcomes depend on marine protected areas with five key features. Nature 506 , 216-220 (2014). https://doi.org/10.1038/nature13022 Dureuil, M., Boerder, K., Burnett, K. A., Froese, R. & Worm, B. Elevated trawling inside protected areas undermines conservation outcomes in a global fishing hot spot. Science 362 , 1403-1407 (2018). https://doi.org/10.1126/science.aau0561 Costello, M. J. & Ballantine, B. Biodiversity conservation should focus on no-take Marine Reserves: 94% of Marine Protected Areas allow fishing. Trends in Ecology & Evolution 30 , 507-509 (2015). https://doi.org/10.1016/j.tree.2015.06.011 Barnes, M. D., Glew, L., Wyborn, C. & Craigie, I. D. Prevent perverse outcomes from global protected area policy. Nature Ecology & Evolution 2 , 759-762 (2018). https://doi.org/10.1038/s41559-018-0501-y Pike, E. P. et al. Ocean protection quality is lagging behind quantity: Applying a scientific framework to assess real marine protected area progress against the 30 by 30 target. Conservation Letters 17 (2024). https://doi.org/10.1111/conl.13020 Council of Europe. Convention on the Conservation of European Wildlife and Natural Habitats. CETS 104, Bern, (1979). European Council. Directive 2009/147/EC of the European Parliament and of the Council on the Conservation of Wild Birds (2009). Centre for International Law. Convention on the Protection of the Black Sea Against Pollution (Bucharest Convention), Bucharest (1992). Raftopoulos, E. The Barcelona Convention System for the Protection of the Mediterranean Sea against Pollution: An international trust at work. Int'l J. Estuarine & Coastal L., 7, p.27. International Journal of Estuarine and Coastal Law 7 , 27-42 (1992). Tromp, D. & Wieriks, K. The OSPAR Convention: 25 years of North Sea protection. Marine Pollution Bulletin 29 , 622-626 (1994). https://doi.org/10.1016/0025-326x(94)90698-x Fitzmaurice, M. The Helsinki Conventions 1974 and 1992. The International Journal of Marine and Coastal Law 13 , 379-394 (1998). Aminian-Biquet, J. et al. Over 80% of the European Union’s marine protected area only marginally regulates human activities. One Earth 7 , 1614-1629 (2024). https://doi.org/10.1016/j.oneear.2024.07.010 European Commission. EU Biodiversity Strategy for 2030: Bringing nature back into our lives. Brussels. Available at: https://eur-lex.europa.eu/legal-content/EN/TXT/?uri=CELEX:52020DC0380 (2020). European Commission. Criteria and guidance for protected areas designations. Brussels. Available at: https://environment.ec.europa.eu/publications/criteria-and-guidance-protected-areas-designations-staff-working-document_en (2022). Horta e Costa, B., Stephenson, F. & Claudet, J. European Union’s strict conservation targets should guide global marine policy. Nature 642 , 38-38 (2025). https://doi.org/10.1038/d41586-025-01731-7 Stephenson, F. et al. Quality of marine protected areas is critical to achieving global biodiversity targets. npj Ocean Sustainability 4 (2025). https://doi.org/10.1038/s44183-025-00169-8 Amengual, J. & Alvarez-Berastegui, D. A critical evaluation of the Aichi Biodiversity Target 11 and the Mediterranean MPA network, two years ahead of its deadline. Biological Conservation 225 , 187-196 (2018). https://doi.org/10.1016/j.biocon.2018.06.032 EEA. State of Europe’s Seas, Technical report No. 2., Copenhagen (2015). Stephenson, F. et al. Implications for the conservation of deep-water corals in the face of multiple stressors: A case study from the New Zealand region. Journal of Environmental Management 346 , 118938 (2023). https://doi.org/10.1016/j.jenvman.2023.118938 Zelli, E. et al. Identifying climate refugia for vulnerable marine ecosystem indicator taxa under future climate change scenarios. Journal of Environmental Management 373 , 122635 (2025). https://doi.org/10.1016/j.jenvman.2024.122635 OBIS. Ocean Biodiversity Information System. Intergovernmental Oceanographic Commission of UNESCO (2026). Assis, J. et al. Bio‐ORACLE v3.0. Pushing marine data layers to the CMIP6 Earth System Models of climate change research. Global Ecology and Biogeography 33 (2024). https://doi.org/10.1111/geb.13813 Principe, S. C. et al. Mapping marine species distributions to inform the design of protected areas in Europe [Manuscript submitted for publication]. Scientific Data (2026). Assis, J., Fragkopoulou, E., Gouvêa, L., Araújo, M. B. & Serrão, E. A. Kelp forest diversity under projected end‐of‐century climate change. Diversity and Distributions 30 (2024). https://doi.org/10.1111/ddi.13837 Gidden, M. J. et al. Global emissions pathways under different socioeconomic scenarios for use in CMIP6: a dataset of harmonized emissions trajectories through the end of the century. Geoscientific Model Development 12 , 1443-1475 (2019). https://doi.org/10.5194/gmd-12-1443-2019 Davies, T. E., Maxwell, S. M., Kaschner, K., Garilao, C. & Ban, N. C. Large marine protected areas represent biodiversity now and under climate change. Scientific Reports 7 (2017). https://doi.org/10.1038/s41598-017-08758-5 Giakoumi, S. et al. Revisiting “Success” and “Failure” of Marine Protected Areas: A Conservation Scientist Perspective. Frontiers in Marine Science 5 (2018). https://doi.org/10.3389/fmars.2018.00223 Blowes, S. A. et al. Mediterranean marine protected areas have higher biodiversity via increased evenness, not abundance. Journal of Applied Ecology 57 , 578-589 (2020). https://doi.org/10.1111/1365-2664.13549 Sullivan-Stack, J. et al. Assessments of expected MPA outcomes can inform and improve biodiversity conservation: Case studies using The MPA Guide. Marine Policy 170 , 106364 (2024). https://doi.org/10.1016/j.marpol.2024.106364 Moilanen, A., Kujala, H. & Leathwick, J. R. The Zonation Framework and Software for Conservation Prioritization. 196-210, Oxford University Press, Oxford (2009). Zhao, Q., Huang, H. & Costello, M. J. Systematic planning shows more than half of the most species-rich ocean region is needed to include all species in representative protected areas. Global Ecology and Conservation 53 , e03036 (2024). https://doi.org/10.1016/j.gecco.2024.e03036 Lauer, D. & Reaka, M. Depth distributions of benthic and pelagic species highlight the potential of mesophotic and deep habitats to serve as marine refugia. Marine Ecology Progress Series 700 , 39-52 (2022). https://doi.org/10.3354/meps14180 Marrocco, V. et al. Behavioural Constraints to Home Range Allometries in Aquatic Organisms. Ecology and Evolution 15 (2025). https://doi.org/10.1002/ece3.71886 Möllmann, C., Müller-Karulis, B., Kornilovs, G. & St John, M. A. Effects of climate and overfishing on zooplankton dynamics and ecosystem structure: regime shifts, trophic cascade, and feedback loops in a simple ecosystem. ICES Journal of Marine Science 65 , 302-310 (2008). https://doi.org/10.1093/icesjms/fsm197 Roberts, K. E., Smith, B. J., Burkholder, D. & Hart, K. M. Evaluating the use of marine protected areas by endangered species: A habitat selection approach. Ecological Solutions and Evidence 2 (2021). https://doi.org/10.1002/2688-8319.12035 Gilmour, M. E. et al. Evaluation of MPA designs that protect highly mobile megafauna now and under climate change scenarios. Global Ecology and Conservation 35 , e02070 (2022). https://doi.org/10.1016/j.gecco.2022.e02070 Predragovic, M. et al. Up to 80% of threatened and commercial species across European marine protected areas face novel climates under high emission scenario. npj Ocean Sustainability 3 (2024). https://doi.org/10.1038/s44183-024-00068-4 Poloczanska, E. S. et al. Global imprint of climate change on marine life. Nature Climate Change 3 , 919-925 (2013). https://doi.org/10.1038/nclimate1958 Burrows, M. T. et al. Ocean community warming responses explained by thermal affinities and temperature gradients. Nature Climate Change 9 , 959-963 (2019). https://doi.org/10.1038/s41558-019-0631-5 Gordó-Vilaseca, C., Stephenson, F., Coll, M., Lavin, C. & Costello, M. J. Three decades of increasing fish biodiversity across the northeast Atlantic and the Arctic Ocean. Proceedings of the National Academy of Sciences 120 (2023). https://doi.org/10.1073/pnas.2120869120 Gordó-Vilaseca, C. et al. Future trends of marine fish biomass distributions from the North Sea to the Barents Sea. Nature Communications 2024 15:1 15 (2024). https://doi.org/10.1038/s41467-024-49911-9 Bartlett, B. S., Erisman, B. & Asch, R. G. Current Marine Protected Areas Conserve Fish Spawning Aggregations Under Climate Change due to Habitat Refugia. Global Change Biology 31 (2025). https://doi.org/10.1111/gcb.70433 Malanoski, C. M., Farnsworth, A., Lunt, D. J., Valdes, P. J. & Saupe, E. E. Climate change is an important predictor of extinction risk on macroevolutionary timescales. Science 383 , 1130-1134 (2024). https://doi.org/10.1126/science.adj5763 Manes, S. et al. Endemism increases species' climate change risk in areas of global biodiversity importance. Biological Conservation 257 , 109070 (2021). https://doi.org/10.1016/j.biocon.2021.109070 Mazaris, A. D. et al. Threats to marine biodiversity in European protected areas. Science of The Total Environment 677 , 418-426 (2019). https://doi.org/10.1016/j.scitotenv.2019.04.333 Oregon State University, IUCN World Commission on Protected Areas - Marine, Marine Conservation Institute, National Geographic Pristine Seas & UN Environment Programme World Conservation Monitoring Centre. The MPA Guide User Manual, version 1., https://mpa-guide.protectedplanet.net (2023). Hermoso, V. et al. The EU Biodiversity Strategy for 2030: Opportunities and challenges on the path towards biodiversity recovery. Environmental Science & Policy 127 , 263-271 (2022). https://doi.org/10.1016/j.envsci.2021.10.028 Gill, D. A. et al. Capacity shortfalls hinder the performance of marine protected areas globally. Nature 543 , 665-669 (2017). https://doi.org/10.1038/nature21708 Horta e Costa, B. et al. Marine protected areas stage of establishment and level of protection are good predictors of their conservation outcomes. Cell Reports Sustainability 2 , 100345 (2025). https://doi.org/10.1016/j.crsus.2025.100345 Zucchetta, M. et al. Can the Effects of Anthropogenic Pressures and Environmental Variability on Nekton Fauna Be Detected in Fishery Data? Insights from the Monitoring of the Artisanal Fishery Within the Venice Lagoon. Estuaries and Coasts 39 , 1164-1182 (2016). https://doi.org/10.1007/s12237-015-0064-y Costello, M. J. Evidence of economic benefits from marine protected areas. Scientia Marina 88 , e080 (2024). https://doi.org/10.3989/scimar.05417.080 Costello, M. J. Fully protected Marine Protected Areas do not displace fisheries. Proceedings of the National Academy of Sciences 121 (2024). https://doi.org/10.1073/pnas.2412543121 European Environment Agency. Marine Protected Areas (MPA) in EEA marine assessment areas, Ver. 2021. In: AGENCY, E. E. (ed.). DOI: eea_v_3035_100_k_mpa-in-marine-assessment-area_p_2021_v01_r00 (2023). Sim, E. Regulations of Human Activities & Protection Levels in Marine Protected Areas of the United Kingdom. [dataset], Figshare (2025). https://doi.org/10.6084/m9.figshare.29901479.v5. Horta e Costa, B., Sousa, I., Kleiven, A. R., Aarflot, J. M. & Moland, E. Data on Marine Protection Levels in Norwegian MPAs: A National Application of The MPA Guide. Data in Brief (submitted). UNEP-WCMC and IUCN. Protected Planet: The World Database on Protected Areas (WDPA) (2026). https://doi.org/10.34892/6fwd-af11. Costello, M. J., Gordó-Vilaseca, C. & Coll, M. Trophic Cascades and Marine Reserves: Dual Indicators of Fishery and Climate Change Disruption in Pelagic and Benthic Ecosystems. In: Imperiled: The Encyclopedia of Conservation . 903-911, Elsevier (2022). Hirzel, A. H., Le Lay, G., Helfer, V., Randin, C. & Guisan, A. Evaluating the ability of habitat suitability models to predict species presences. Ecological Modelling 199 , 142-152 (2006). https://doi.org/10.1016/j.ecolmodel.2006.05.017 Costello, M. J. et al. Marine biogeographic realms and species endemicity. Nature Communications 8 (2017). https://doi.org/10.1038/s41467-017-01121-2 Liu, C., Newell, G. & White, M. On the selection of thresholds for predicting species occurrence with presence‐only data. Ecology and Evolution 6 , 337-348 (2016). https://doi.org/10.1002/ece3.1878 Liu, C., White, M. & Newell, G. Selecting thresholds for the prediction of species occurrence with presence‐only data. Journal of Biogeography 40 , 778-789 (2013). https://doi.org/10.1111/jbi.12058 Boettiger, C., Lang, D. T. & Wainwright, P. C. rfishbase: exploring, manipulating and visualizing FishBase data from R. Journal of Fish Biology 81 , 2030-2039 (2012). https://doi.org/10.1111/j.1095-8649.2012.03464.x Principe, S. et al. MPA Europe Deliverable 3.3. Database of species and habitats conservation status. Zenodo (2024). https://doi.org/10.5281/zenodo.11075390 Principe, S. et al. Publish maps and models of biogenic habitat distribution in Europe on EMODnet. Zenodo (2023). https://doi.org/10.5281/zenodo.10422129 R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, Vienna, Austria, 2025). https://www.R-project.org/. Hijmans. terra: Spatial Data Analysis. R package version 1.8-93. https://cran.r-project.org/web/packages/terra/index.html (2026). Baston. exactextractr: Fast Extraction from Raster Datasets using Polygons. R package version 0.10.0 https://CRAN.R-project.org/package=exactextractr (2023). Additional Declarations There is NO Competing Interest. Supplementary Files EskucheKeithetalsupplementary.docx European Marine Protected Areas are well-placed and climate-resilient, but weakly protected - Supplementary material Cite Share Download PDF Status: Under Review Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8894972","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":605762727,"identity":"185d3f4b-2e0d-4da8-b942-ab06cb8d006f","order_by":0,"name":"Fabrice Stephenson","email":"data:image/png;base64,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","orcid":"","institution":"School of Natural and Environmental Sciences, Newcastle University; IUCN World Commission on Protected Areas","correspondingAuthor":true,"prefix":"","firstName":"Fabrice","middleName":"","lastName":"Stephenson","suffix":""},{"id":605762728,"identity":"3079bbf0-ec30-4e08-98f9-e03d054ca019","order_by":1,"name":"Patrick Eskuche-Keith","email":"","orcid":"","institution":"School of Natural and Environmental Sciences, Newcastle University; IUCN World Commission on Protected Areas,","correspondingAuthor":false,"prefix":"","firstName":"Patrick","middleName":"","lastName":"Eskuche-Keith","suffix":""},{"id":605762729,"identity":"0a3931b4-6171-4bf6-a0ab-f582291a734b","order_by":2,"name":"Anna Addamo","email":"","orcid":"https://orcid.org/0000-0002-1228-2143","institution":"Faculty of Biosciences and Aquaculture, Nord University","correspondingAuthor":false,"prefix":"","firstName":"Anna","middleName":"","lastName":"Addamo","suffix":""},{"id":605762730,"identity":"3b90db86-0f0c-4006-bac9-ae4cd45fae9b","order_by":3,"name":"Peter Almond","email":"","orcid":"https://orcid.org/0009-0003-3419-9169","institution":"School of Natural and Environmental Sciences, Newcastle University","correspondingAuthor":false,"prefix":"","firstName":"Peter","middleName":"","lastName":"Almond","suffix":""},{"id":605762731,"identity":"003bc321-c973-4b2f-8b0e-d4bcd2849871","order_by":4,"name":"Barbara Horta e Costa","email":"","orcid":"https://orcid.org/0000-0002-9960-3893","institution":"CCMAR - Centre of Marine Sciences of the Algarve","correspondingAuthor":false,"prefix":"","firstName":"Barbara","middleName":"Horta e","lastName":"Costa","suffix":""},{"id":605762732,"identity":"85e707bd-53f3-46f2-a965-a91fb3e8a79e","order_by":5,"name":"Silas Principe","email":"","orcid":"","institution":"Ocean Biodiversity Information System, International Oceanographic Data and Information Exchange, Intergovernmental Oceanographic Commission, UNESCO","correspondingAuthor":false,"prefix":"","firstName":"Silas","middleName":"","lastName":"Principe","suffix":""},{"id":605762733,"identity":"0c43c93a-f07e-41ea-bb88-f5aa931cdfe4","order_by":6,"name":"Jorge Assis","email":"","orcid":"https://orcid.org/0000-0002-6624-4820","institution":"Centre of Marine Sciences, University of Algarve, Faro, Portugal","correspondingAuthor":false,"prefix":"","firstName":"Jorge","middleName":"","lastName":"Assis","suffix":""},{"id":605762734,"identity":"1003e28b-a2fd-4b29-a4f6-3f5fb2a578a3","order_by":7,"name":"Inês Sousa","email":"","orcid":"https://orcid.org/0000-0002-2638-0094","institution":"Centre of Marine Sciences (CCMAR/CIMAR LA), Universidade do Algarve","correspondingAuthor":false,"prefix":"","firstName":"Inês","middleName":"","lastName":"Sousa","suffix":""},{"id":605762735,"identity":"4c6b6df5-eca0-4788-8494-acd48eee842b","order_by":8,"name":"Even Moland","email":"","orcid":"","institution":"Institute of Marine Research; Centre for Coastal Research, University of Agder","correspondingAuthor":false,"prefix":"","firstName":"Even","middleName":"","lastName":"Moland","suffix":""},{"id":605762736,"identity":"1aee6f02-9e64-4925-9836-51475ddeecbe","order_by":9,"name":"Alf Kleiven","email":"","orcid":"","institution":"Institute of Marine Research, Norway","correspondingAuthor":false,"prefix":"","firstName":"Alf","middleName":"","lastName":"Kleiven","suffix":""},{"id":605762737,"identity":"e4d0d351-c91b-466f-b134-b5252ec943b7","order_by":10,"name":"Johanna Aarflot","email":"","orcid":"https://orcid.org/0000-0003-2481-0190","institution":"Institute of Marine Research, Norway","correspondingAuthor":false,"prefix":"","firstName":"Johanna","middleName":"","lastName":"Aarflot","suffix":""},{"id":605762738,"identity":"423f946e-40e8-417b-8243-20af5d0c5560","order_by":11,"name":"Pippa Moore","email":"","orcid":"","institution":"New Castle University","correspondingAuthor":false,"prefix":"","firstName":"Pippa","middleName":"","lastName":"Moore","suffix":""},{"id":605762739,"identity":"16bdec76-7d30-4d99-a296-e9a32244c360","order_by":12,"name":"Daniel Skerritt","email":"","orcid":"","institution":"School of Natural and Environmental Sciences, Newcastle University; Transparent Oceans Initiative, Oceana","correspondingAuthor":false,"prefix":"","firstName":"Daniel","middleName":"","lastName":"Skerritt","suffix":""},{"id":605762741,"identity":"dc621f47-0342-47ad-92bc-adaea97bd34a","order_by":13,"name":"Elsa Sim","email":"","orcid":"","institution":"Faculty of Biosciences and Aquaculture, Nord University","correspondingAuthor":false,"prefix":"","firstName":"Elsa","middleName":"","lastName":"Sim","suffix":""},{"id":605762743,"identity":"33e4db1a-fec7-4f3f-a6b9-be6e7b152ba8","order_by":14,"name":"Mark Costello","email":"","orcid":"https://orcid.org/0000-0003-2362-0328","institution":"Nord University","correspondingAuthor":false,"prefix":"","firstName":"Mark","middleName":"","lastName":"Costello","suffix":""}],"badges":[],"createdAt":"2026-02-16 16:56:28","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8894972/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8894972/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":106724658,"identity":"440a8ec1-90c8-43f4-8860-7cd7eedb122b","added_by":"auto","created_at":"2026-04-12 18:29:01","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1968973,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of marine protected areas (MPAs) across the study area, coloured by protection level (adapted and updated from \u003csup\u003e25\u003c/sup\u003e). Point size indicates the relative area of each MPA. The study area was split into multiple subregions based on a combination of the International Council for the Exploration of the Sea (ICES) ecoregions (for the North Sea, Celtic Seas, Norwegian Sea, Icelandic and Faroese waters, and the Barents Sea and Arctic Ocean), and Marine Strategy Framework Directive for all other subregions. Donut plots indicate the proportion of each MPA protection level relative to the total area covered by MPAs in each subregion, with values in the centre of each donut indicating the total proportion of each subregion covered by MPAs. The large donut in the top left reflects the MPA coverage of the entire pan-European study area. Coverage proportions below 0.1% are not highlighted with text. Light grey shading and dashed lines indicate national Exclusive Economic Zones (EEZs), representing the extent of the study area.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/bfd311efcab91a37019420c2.png"},{"id":106495276,"identity":"fb734434-b3e8-4c4c-a015-a296ffac2719","added_by":"auto","created_at":"2026-04-09 08:13:48","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1015297,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage of species distribution overlaps with marine protected area (MPA) protection levels, by biotic group. Values on the x-axis represent the proportion of species ranges covered by each MPA protection level as a percentage of their total distribution area in pan-European seas. Protection levels of Fully and Highly are considered to provide strong protection, while all other protection levels (excluding Unclassified) represent weak protection. The x-axis has been cut to omit the top 5 % of values in each MPA protection level for clarity (but these datapoints are still incorporated in the definition of the boxplots: vertical bars indicate the 25 % quantile, median and 75 % quantile of the data; whiskers indicate the full range excluding outliers). Numbers in brackets after MPA protection levels on the y-axis indicate the area of each as a percentage of the total area of pan-European seas, while \u003cem\u003en \u003c/em\u003evalues indicate the total number of species included in each biotic group.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/0eec7bbbf39e6903f9abc5b1.png"},{"id":106724532,"identity":"bf3c068a-4d3b-4dee-a27c-f62a97a24217","added_by":"auto","created_at":"2026-04-12 18:28:30","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":548666,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage of species distribution overlaps with marine protected area (MPA) protection levels, for (a) threatened species (IUCN Red List Vulnerable, Endangered or Critically endangered species) and (b) habitat-forming species (biogenic reefs, canopy-forming macroalgae and seagrasses). Values on the x-axis represent the proportion of species ranges covered by each MPA protection level as a percentage of their total distribution area in pan-European seas. The x-axis has been cut to omit the top 5 % percentile values in each MPA protection level for clarity (but these datapoints are still incorporated in the boxplots: vertical bars show the 25 % quantile, median and 75 % quantile of the data; whiskers show the full range excluding outliers). Numbers in brackets after MPA protection levels on the y-axis indicate the area of each as a percentage of the total area of pan-European seas, while \u003cem\u003en\u003c/em\u003evalues refer to the total number of species included in each panel.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/2433d377843198d6aed9f743.png"},{"id":106495143,"identity":"bd84d533-74e6-408d-be22-24703a1050fe","added_by":"auto","created_at":"2026-04-09 08:13:21","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1148333,"visible":true,"origin":"","legend":"\u003cp\u003eRelative change in overlap between species ranges and marine protected areas (MPAs) by biotic group under the intermediate Shared Socio-economic Pathway emissions scenario (SSP2-4.5), for taxa classified as ‘High climate-risk’. Values beyond the range of the 2.5 % and 97.5 % quantiles of the data are omitted for clarity. Circles show the median value across the full data range. Percentages on either side of the whiskers identify the proportion of species for each combination of biotic group and MPA protection level which are projected to experience either increases or decreases in overlap (note that these values do not always sum to 100% as they do not consider species with zero change in coverage). The \u003cem\u003en\u003c/em\u003e values indicate the total number of species included in each panel.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/c034ac4e01b2d34c5e101b32.png"},{"id":106726078,"identity":"52cf9171-4a97-4086-92f9-da44f88f4dbe","added_by":"auto","created_at":"2026-04-12 18:35:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5838665,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/688fa06d-ddd8-4238-bf6c-904325b7721a.pdf"},{"id":106495242,"identity":"5b5b5352-a1f7-4706-be02-efc5fdb201c8","added_by":"auto","created_at":"2026-04-09 08:13:30","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":806542,"visible":true,"origin":"","legend":"European Marine Protected Areas are well-placed and climate-resilient, but weakly protected - Supplementary material","description":"","filename":"EskucheKeithetalsupplementary.docx","url":"https://assets-eu.researchsquare.com/files/rs-8894972/v1/6e7034e4aae6b18749e4698c.docx"}],"financialInterests":"There is \u003cb\u003eNO\u003c/b\u003e Competing Interest.","formattedTitle":"European Marine Protected Areas are well-placed and climate-resilient, but weakly protected","fulltext":[{"header":"Introduction","content":"\u003cp\u003eGlobally, marine biodiversity is declining at an unprecedented rate, driven primarily by human activities that further degrade habitats, pollute ecosystems, and intensify resource extraction \u003csup\u003e1,2\u003c/sup\u003e. Climate change adds to these pressures by raising ocean temperature and consequently changing species distributions, with thousands of marine species having already shifted their distribution poleward \u003csup\u003e3,4\u003c/sup\u003e. Together, these drivers transform marine ecosystems at rates frequently exceeding the practical capacity of existing management systems.\u003c/p\u003e\n\u003cp\u003eMarine Protected Areas (MPAs) are effective tools for reducing cumulative stressors, safeguarding biodiversity, and sustaining ecosystem services \u003csup\u003e5-7\u003c/sup\u003e. By protecting locations likely to remain suitable under future climates, MPAs can support ecological resilience \u003csup\u003e8\u003c/sup\u003e. These attributes underpin global conservation commitments, including targets of the Global Biodiversity Framework and the recent High Seas Treaty \u003csup\u003e9\u003c/sup\u003e. However, expansion of MPA networks alone does not guarantee biodiversity protection or climate readiness; effectiveness depends on placement, restricted pressures, enforcement, and stakeholder participation \u003csup\u003e10\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003eDespite growing global MPA coverage, evidence suggests that current networks insufficiently represent and protect marine biodiversity. Only a minority of species have substantial range fractions within existing MPAs \u003csup\u003e11-13\u003c/sup\u003e, and most designated sites continue to allow ecologically damaging activities, such as industrial fishing including seabed trawling \u003csup\u003e14,15\u003c/sup\u003e, eroding intended conservation outcomes \u003csup\u003e16,17\u003c/sup\u003e. To address inconsistent definitions and expectations of protection, the MPA Guide provides a framework linking protection levels to expected ecological outcomes \u003csup\u003e6\u003c/sup\u003e. Its global application revealed that only one-third of the 100 largest MPAs (90 % of global MPA coverage, representing 7.3 % of global ocean area) qualify as \u0026lsquo;Fully\u0026rsquo; or \u0026lsquo;Highly\u0026rsquo; protected \u003csup\u003e18\u003c/sup\u003e, underscoring the disparity between nominal protection and realised ecological security (the so-called \u0026lsquo;paper parks\u0026rsquo; phenomenon).\u003c/p\u003e\n\u003cp\u003eAcross Europe, conservation ambition is shaped both by global commitments and also by a complex patchwork of regional and national governance frameworks. European Union (EU) legislation, notably the Birds and Habitats Directives \u003csup\u003e19,20\u003c/sup\u003e, supports the Natura 2000 network, while regional conventions (e.g., OSPAR, the Helsinki Convention, the Bucharest Convention and the Barcelona Convention) complement EU policy across basin scales \u003csup\u003e21-24\u003c/sup\u003e. However, these frameworks historically emphasise spatial designation rather than ecological representativity or specific management requirements, producing a heterogeneous mosaic of MPA types and regulatory regimes. This complexity complicates assessments of true protection levels and coordination of coherent, climate-ready conservation across jurisdictions.\u003c/p\u003e\n\u003cp\u003eWithin EU waters, MPAs cover \u0026gt;12 % of marine areas, yet only 1.5 % is classified as \u0026lsquo;strongly protected\u0026rsquo; (i.e., falling within the \u0026lsquo;Fully\u0026rsquo; or \u0026lsquo;Highly\u0026rsquo; protected categories under the MPA Guide) \u003csup\u003e6,25\u003c/sup\u003e. The EU\u0026rsquo;s Biodiversity Strategy for 2030 aims to designate 30 % of Earth\u0026rsquo;s ocean area as protected areas by 2030 (\u0026ldquo;30 \u0026times; 30\u0026rdquo;) and ensure at least 10 % of European seas are \u0026lsquo;strictly\u0026rsquo; protected (i.e., left essentially undisturbed from human pressures and threats, corresponding to full protection as defined in the MPA guide) \u003csup\u003e6,26,27\u003c/sup\u003e. The EU\u0026rsquo;s ambition will require both substantial expansion and the effective management of MPAs capable of delivering strong ecological outcomes \u003csup\u003e28,29\u003c/sup\u003e. Critically, assessments of protection levels have not extended to non-EU European marine nations, including Iceland, Norway, the Faroe Islands, and the United Kingdom, which collectively manage a large share of pan-European seas and are ecologically connected. Consequently, the actual extent of marine protection in Europe remains unclear.\u003c/p\u003e\n\u003cp\u003eA central uncertainty is whether existing MPA locations adequately capture marine species\u0026rsquo; distributions, both under present-day conditions and as ranges shift under climate change. As species ranges expand or contract due to shifting conditions \u003csup\u003e30\u003c/sup\u003e, increasing proportions of their ranges may fall outside or inside current MPA boundaries, raising the question of the adequacy of existing networks \u003csup\u003e31-33\u003c/sup\u003e. Recent advances in open-access biodiversity and environmental datasets, such as OBIS \u003csup\u003e34\u003c/sup\u003e and Bio-ORACLE v3 \u003csup\u003e35\u003c/sup\u003e, together with high-resolution species distribution models (SDMs) for thousands of marine taxa \u003csup\u003e36\u003c/sup\u003e, enable forecasts of future species distributions under alternative emissions scenarios \u003csup\u003e37\u003c/sup\u003e. These developments offer opportunities to evaluate species coverage by MPAs under climate change.\u003c/p\u003e\n\u003cp\u003eHere we assessed how MPAs with protection levels classified using the MPA Guide overlapped with predicted distributions for 9,794 marine species across pan-European seas; defined as the Exclusive Economic Zones of EU member states, Iceland, Norway (including Svalbard and Jan Mayen), the Faroe Islands, and the United Kingdom. We quantified the proportion of current species distributions encompassed by MPAs across protection levels and major biotic groups, and evaluated climate readiness by comparing present-day distributions with those projected for the end of the century (2090-2100) under a moderate \u0026lsquo;most likely\u0026rsquo; Shared Socio-economic Pathway emissions scenario (SSP2-4.5) \u003csup\u003e38\u003c/sup\u003e. We categorised species into climate-risk classes based solely on projected changes in range size (e.g.\u0026nbsp;\u003csup\u003e32\u003c/sup\u003e), whereby \u0026lsquo;High climate-risk\u0026rsquo; species were projected to lose \u0026gt; 50 % of their present-day distribution, \u0026lsquo;Medium climate-risk\u0026rsquo; species were projected to lose 10-50 % of their distribution, and \u0026lsquo;Low climate-risk\u0026rsquo; species were projected to lose \u0026lt;10 % or even gain distribution area. We examined whether MPAs were positioned to protect these classes as climatic conditions change. Through this analysis, we provide the first pan-European assessment of the effectiveness and climate readiness of current marine protection across all European jurisdictions.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003eDistribution of MPAs by protection level\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eMPAs covered 1,017,565 km\u0026sup2;, approximately 11 % of pan-European seas. Most (91 %) offered weak protection (i.e., \u0026lsquo;Light\u0026rsquo;, \u0026rsquo;Minimal\u0026rsquo; or \u0026lsquo;Incompatible\u0026rsquo;), while only ~1 % were strongly protected (i.e., \u0026lsquo;Fully\u0026rsquo; or \u0026lsquo;Highly\u0026rsquo;), with the remainder unclassified (Figure 1, Table 1). Individual national Exclusive Economic Zones (EEZs) showed considerable variability, with MPA coverage ranging from \u0026lt;1 % to ~48 %, although all countries had \u0026lt;1 % of their EEZ under strong protection (Table S1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCoverage of species ranges by MPA protection level\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAcross all species, median MPA coverage of distributions was 25 % (Figure 2, Table S2). Among the 4,917 species with \u0026ge; 25 % of their ranges falling within MPA boundaries (i.e., \u0026ge; overall median), median coverage by strongly protected MPAs was \u0026lt; 1 % (95th percentile = 5 %). Overall, across all species, strongly protected MPAs generally covered \u0026lt; 1 % of species\u0026rsquo; distributions (median \u0026lt;1%), whereas weakly protected MPAs accounted for most coverage (median = 22 %; Table S2).\u003c/p\u003e\n\u003cp\u003eAcross biotic groups, median coverage of species distributions by all MPAs ranged from 14 % (pelagic invertebrates) to 30 % (demersal fish) (Figure 2, Table S2). Median coverage by MPAs with strong protection levels remained \u0026lt; 1 % for all groups, with 95th percentiles \u0026lt; 2 % in all cases (Table S2). Coverage by weak protection levels ranged from 12 % (pelagic fish and invertebrates) to 25 % (benthic invertebrates).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eMedian coverage of threatened and habitat-forming species distributions across all MPAs was 29 % and 25 %, respectively (Table S3). Among threatened species (i.e., IUCN Red List Vulnerable, Endangered or Critically endangered species), two thirds (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 146) had \u0026ge; 25 % of their individual species\u0026rsquo; distribution protected. For habitat-formers (biogenic reefs, canopy-forming macroalgae and seagrasses), half of the species (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 62) met the threshold of \u0026ge; 25 % protection. Median coverage by strongly protected MPAs was \u0026lt; 1 % for both threatened and habitat-forming species, while weakly protected MPAs accounted for substantially higher coverage (26 % and 22 %, respectively) (Figure 3, Table S3).\u003c/p\u003e\n\u003cp\u003eTable 1: Total areas (km\u003csup\u003e2\u003c/sup\u003e) covered by each MPA protection level. Values in brackets express this as a percentage of the total area covered by MPAs in each column. Combined areas are presented for the Icelandic, Faroese, Norwegian and UK assessments and also for the full pan-European study region. For the Icelandic and Faroese MPAs, a lack of sufficient information for assessment meant these were recorded as Unclassified. The area of each protection level as a percentage of the total area of pan-European seas is also presented.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"602\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 112px;\"\u003e\n \u003cp\u003eCombined protection level\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eMPA Guide protection level\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003eUK, Norway, Iceland and Faroes MPAs\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003epan-European study region\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003ePercentage of pan-European seas (%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" valign=\"top\" style=\"width: 112px;\"\u003e\n \u003cp\u003eStrong\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eFully\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e57 (0.1)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1,156 (0.1)\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e\u0026lt; 0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eHighly\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e1,003 (0.2)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e8,339 (0.8)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e0.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"3\" valign=\"top\" style=\"width: 112px;\"\u003e\n \u003cp\u003eWeak\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLightly\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e24,507 (6.0)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e110,737 (10.0)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e1.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eMinimally\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e170,850 (41.5)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e473,584 (46.5)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e5.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eIncompatible\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e221,746 (51.5)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e342,543 (33.7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 112px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eUnclassified\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e3,361 (0.8)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e81,207 (8.0)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 112px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eTOTAL\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 141px;\"\u003e\n \u003cp\u003e411,525\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1,017,565\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 132px;\"\u003e\n \u003cp\u003e11.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003cem\u003eImpacts of climate change on species distributions\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eA marginally smaller subset of spatial predictions (\u003cem\u003en\u003c/em\u003e = 9,635) was available under both present-day environmental conditions and the SSP2-4.5 scenario, due to filtering of SDM outputs during the data processing stages (see Methods). These were used to investigate potential future range changes. Approximately 54 % of these species (\u003cem\u003en\u003c/em\u003e = 5,227) were projected to experience net decreases in their distribution area within pan-European waters, while ~45 % (\u003cem\u003en\u003c/em\u003e = 4,357) were projected to experience net increases and 51 species were not projected to show a net area change.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThere was high variability in projected changes in distribution area across biotic groups. Primary producers and both pelagic and benthic invertebrates were dominated by species projected to experience decreases in distribution area within the study region, with median losses of 7 %, 6 % and 3 %, respectively. In contrast, the majority of elasmobranchs were projected to experience distribution area increases, with median gains of 7 %, while mammals and demersal and pelagic fish were projected to experience median increases of ~ 2 % overall. \u0026lsquo;High climate-risk\u0026rsquo; species (i.e., defined in this study as those projected to lose \u0026gt; 50 % of their present-day distribution area under climate change) comprised ~18 % of taxa (\u003cem\u003en\u003c/em\u003e = 1,723), while \u0026lsquo;Medium climate-risk\u0026rsquo; species (10-50 % loss) accounted for ~25 % (\u003cem\u003en\u003c/em\u003e = 2,404). The remaining ~57 % of taxa (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 5,508) were classed as at \u0026lsquo;Low climate-risk\u0026rsquo; (\u0026lt; 10 % loss or net gains).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eImpacts of climate change on coverage of species distributions by MPA protection levels\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWhen all species were aggregated, the median projected changes in range coverage from MPA protection levels by 2100 under SSP2-4.5 were negligible (\u0026lt; 1 % in either direction) (Table S4). Similarly, when split by biotic group, median change across all MPAs ranged from -1 % (elasmobranchs) to +1 % (mammals) (Table S4). However, changes in coverage varied widely across risk classes and among groups.\u003c/p\u003e\n\u003cp\u003eFor \u0026lsquo;High climate-risk\u0026rsquo; species, median relative change in MPA coverage under SSP2-4.5 was positive across all biotic groups, with increases between \u0026lt; 1% (demersal fish and invertebrates) and ~12 % (mammals) (Figure 4, Table S5). Median changes in the proportion of distributions covered by strongly protected MPAs were minor (\u0026lt; 1 % gains) for all groups while median changes in coverage within weakly protected MPAs ranged from \u0026lt; -1% (demersal fish) to +11 % (mammals) (Table S5).\u003c/p\u003e\n\u003cp\u003eFor \u0026lsquo;Medium climate-risk\u0026rsquo; taxa, the majority of biotic groups were projected to gain proportional coverage by MPAs, up to ~3 % (mammals), although benthic invertebrates were generally projected to lose coverage (median: -1%) (Figure S1, Table S6). Median changes in coverage by strongly protected MPAs were minor (\u0026lt; 1 % in either direction), while median changes in coverage by weakly protected MPAs ranged from \u0026lt; -1 % (benthic invertebrates) to +3 % (mammals) (Table S6).\u003c/p\u003e\n\u003cp\u003eFor \u0026lsquo;Low climate-risk\u0026rsquo; taxa, most species in all groups were projected to experience limited decreases in the proportion of their distributions covered by MPAs (Figure S2, Table S7). Median changes were generally negative but minor, with coverage changes by both strongly and weakly protected MPAs ranging from -1 % to no change (Table S7).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWe provide the first pan-European evaluation of MPA effectiveness integrating predicted distributions for 9,794 marine species with up-to-date MPA protection level classifications. Despite occupying only ~11 % of pan-European seas, we show that MPAs generally encompass one-quarter of species\u0026rsquo; distributions, suggesting they are broadly well placed to protect biodiversity. Encouragingly, these conservation benefits persist under future climate conditions; median coverage changes are negligible (\u0026lt; 1 % across taxa), suggesting the current spatial configuration of MPAs is robust to projected range shifts under an intermediate emissions scenario (SSP2-4.5). However, strong protection remains extremely limited, with \u0026lt; 1 % of species\u0026rsquo; distributions falling within Fully or Highly protected areas. Together, these findings highlight the potential of Europe\u0026rsquo;s MPA distribution and the urgent need to strengthen protection levels to secure biodiversity outcomes now and in the future. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMPAs are well-placed, but not optimised\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe large proportion of species\u0026rsquo; ranges within MPAs relative to their areal footprint indicates their preferential placement in biodiversity-rich regions. This may reflect deliberate prioritisation of ecologically important areas or the concentration of many early designations in high-richness coastal zones \u003csup\u003e39\u003c/sup\u003e. Such patterns are consistent with the idea that spatial planning efforts, whether explicit or emergent, have captured areas of relatively high conservation value \u003csup\u003e40,41\u003c/sup\u003e. Because existing MPAs already overlap substantial parts of many species\u0026rsquo; distributions, strengthening protection and management within current boundaries, particularly in sites with high biodiversity representation, could deliver disproportionately strong conservation outcomes without relying solely on further spatial expansion of MPAs \u003csup\u003e6,42\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSystematic conservation planning has repeatedly shown that alternative configurations can achieve similar or higher representation at lower area cost, including in global analyses using tools such as Zonation \u003csup\u003e13,43,44\u003c/sup\u003e. Our results highlight that while pan-European MPAs are generally well placed (although weakly protected), there remains scope to refine planning for more even representation across taxa and marine regions. For example, pelagic invertebrates and fishes have lower median MPA coverage (14 % and 15 %, respectively) than other groups and substantially lower than the overall median (25 %). This likely reflects these species often having larger ranges than their benthic and demersal counterparts \u003csup\u003e45,46\u003c/sup\u003e. In some cases, this underrepresentation may be less problematic; zooplankton, for instance, are not typically subject to direct fishing pressure, so lower MPA coverage may not equate to major gaps in protection \u003csup\u003e47\u003c/sup\u003e. This may also apply to highly migratory species including certain elasmobranchs or marine mammals, where protecting strategic places (e.g. spawning, nursery and feeding grounds) provides greater benefits than simply increasing overall overlap with MPAs \u003csup\u003e48,49\u003c/sup\u003e. By contrast, for groups exposed to intense anthropogenic impacts, such as many widespread pelagic fishes or benthic invertebrates sensitive to bottom-contact fishing, low representation within MPAs could indicate missed opportunities for targeted improvements in spatial coverage and management \u003csup\u003e50\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eClimate-driven change and the resilience of spatial placement\u003c/em\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eProjected changes in species distributions reveal some climate-driven redistribution across pan-European seas by the end of the century. Under SSP2-4.5, approximately 54 % of species are expected to shrink their geographic range within European waters, with ~18 % classified here as \u0026lsquo;High climate-risk\u0026rsquo; due to declines in distribution area exceeding 50 %. These projections are specific to the pan-European seas, yet in many cases likely reflect broader poleward or basin-scale redistributions under warming \u003csup\u003e51-54\u003c/sup\u003e. Range changes are uneven across functional groups, with invertebrates and primary producers projected to display median relative losses, whereas marine mammals, fish and elasmobranchs are projected to experience relative gains.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eDespite projected range shifts, the proportion of species\u0026rsquo; distributions falling inside MPAs remains remarkably stable in future. Across all biotic groups and protection levels, median changes in MPA coverage are \u0026lt; 1 %, indicating that current MPAs continue to intersect substantial portions of species\u0026rsquo; distributions, at least under an intermediate emissions scenario. \u0026lsquo;High climate-risk\u0026rsquo; species show a median increase in their representation within MPAs despite losing large fractions of their total pan-European range, implying that most projected loss occurs outside protected areas while currently suitable habitat within MPAs persists. Conversely, \u0026lsquo;Low climate-risk\u0026rsquo; species, particularly those projected to expand their distributions, tend to experience declining representation, reflecting that most MPAs were not designed with future range expansions or poleward shifts in mind \u003csup\u003e10\u003c/sup\u003e. Together, these patterns suggest pan-European MPA placement exhibits substantial climate robustness, with protected areas acting as refugia for species undergoing regional declines, consistent with observations from other marine systems \u003csup\u003e55\u003c/sup\u003e. Our measure of \u0026lsquo;climate-risk\u0026rsquo; provides a simple indicator which could be used to prioritise certain taxa based on their expected range changes. In addition to proportional area loss however, more detailed case studies and assessments should account for overall range size as this is a major predictor of extinction risk under climate change \u003csup\u003e56\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003eThese findings resonate with broader work linking spatial planning to climate-resilient conservation. Designing MPAs solely around present-day habitat suitability is not guaranteed to safeguard species under climate change, whereas integrating projected distributions may improve protection of key climate refugia \u003csup\u003e32\u003c/sup\u003e. In some large MPAs, species representation is projected to increase under climate change, underscoring that existing protections can remain relevant as conditions shift \u003csup\u003e39\u003c/sup\u003e. However, threatened and commercially important species in pan-European MPAs face changing climate conditions, particularly under high-emission scenarios and in semi-enclosed seas such as the Mediterranean, Baltic and Black seas \u003csup\u003e50,57\u003c/sup\u003e. In these regions, even dense MPA networks may struggle to offset climate-driven changes in environmental suitability, unless protection levels are sufficient to safeguard biodiversity \u003csup\u003e42\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eProtection quality is the critical bottleneck\u003c/em\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe most pervasive weakness of present European MPAs lies not in their location but in how strongly they are protected. Across all taxa, fewer than 1 % of species\u0026rsquo; distributions occur within strongly protected MPAs. This means that even species with extensive spatial overlap with MPAs are subject to weak restrictions on extractive activities and other pressures.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThese results support the conclusion that management shortfalls and low protection levels, rather than spatial gaps, are the primary barriers to delivering the biodiversity outcomes promised by MPA policy \u003csup\u003e5,6,16,25,58\u003c/sup\u003e. The poor representation of strongly protected MPAs within all individual EEZs highlights that this is a widespread issue which all relevant European nations need to take legislative action on in order to truly safeguard their marine biodiversity. At the regional scale, improving MPA protection levels would also provide a strategic contribution towards achieving the European 10 % target for strict protection (corresponding to full protection as defined in the MPA guide)\u0026nbsp;\u003csup\u003e6,59,60\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eOur analysis considered protection levels but not the stage of establishment, which can be a critical determinant of ecological performance \u003csup\u003e6\u003c/sup\u003e. Implementation and active management are key components of the MPA Guide and can be critical for MPA effectiveness \u003csup\u003e6,61\u003c/sup\u003e. Actively managed MPAs deliver significantly stronger ecological benefits than sites that remain partially implemented or, worse, are only designated on paper \u003csup\u003e62\u003c/sup\u003e. Thus, increasing the area considered as Fully and/or Highly protected will be necessary but not sufficient without full implementation, enforcement, and adequate resources for management. However, the overwhelming majority of MPAs within pan-European seas have low designated levels of protection, which, even if implemented or actively managed, may provide no measurable ecological benefits and could even have negative impacts (as measured using indicators such as fish density or biomass) \u003csup\u003e62\u003c/sup\u003e. It is therefore imperative that efforts to improve the effectiveness of existing pan-European MPAs focus on both their active management and the prohibition of harmful or destructive activities within their boundaries.\u003c/p\u003e\n\u003cp\u003eThe interaction between climate risk and protection levels is particularly relevant. \u0026lsquo;High climate-risk\u0026rsquo; species are projected to maintain or increase the proportion of their range within MPAs under future conditions, but their low coverage by strong protection undermines the potential benefits of this favourable spatial configuration \u003csup\u003e50\u003c/sup\u003e. Variation among taxa further emphasises the need to align protection levels with ecological need. Groups with limited exposure to direct anthropogenic impacts, such as pelagic invertebrates, may not require extensive strong protection \u003csup\u003e63\u003c/sup\u003e. By contrast, benthic invertebrates, elasmobranchs, and commercially targeted fishes face intense fishing pressure yet receive disproportionately weak protection \u003csup\u003e25,50\u003c/sup\u003e. For these taxa, the scarcity of fully or highly protected areas directly compromises prospects for recovery, resilience, and long-term viability \u003csup\u003e51\u003c/sup\u003e. Taken together, our findings indicate that simply increasing the areal extent of MPAs will not address the core challenges facing marine biodiversity. Expansion alone cannot compensate for inadequate protection levels \u003csup\u003e16,29,62\u003c/sup\u003e or limited management enforcement \u003csup\u003e6,42\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eLimitations and future directions\u003c/em\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eOur analysis is subject to several limitations that should inform interpretation and future work. First, we relied on species distribution models derived from available biodiversity and environmental data; these models inevitably carry uncertainty related to limited and uneven sampling across species\u0026rsquo; geographic ranges, choices of model algorithms and parameters, and accuracy of climate projections \u003csup\u003e36,37\u003c/sup\u003e. Second, we focused on a single intermediate emissions scenario (SSP2-4.5) representing a \u0026lsquo;most-likely\u0026rsquo; future under current global socio-economic trajectories. Higher-emission pathways are likely to produce greater range extensions and contractions in response to changing climate, particularly in enclosed and high-latitude seas \u003csup\u003e54\u003c/sup\u003e. Addressing these points can be accommodated by monitoring actual changes in species distribution and abundance and adapting conservation accordingly. Our analyses were restricted to EEZs and did not consider areas beyond national jurisdiction (ABNJ), which are increasingly important in European and global conservation strategies. With the recent implementation of the High Seas Treaty, there is a clear basis for expanding similar analyses to ABNJ to support systematic spatial prioritisation and conservation planning there.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur results indicate that strengthening protection within the existing MPA network (alongside carefully planned expansion) offers one of the most direct and effective pathways for Europe to meet its marine biodiversity commitments. Major conservation benefits could be realised by upgrading the protection level and ensuring active management of current sites, particularly those that already capture a large share of species\u0026rsquo; distributions or act as climate refugia, while using new designations to fill residual spatial and taxonomic gaps. In this context, the drive to achieve \u0026ldquo;30 \u0026times; 30\u0026rdquo; should not only increase the areal extent of MPAs, but must prioritise Fully and Highly protected areas and ensure that all MPAs are effectively managed, ecologically coherent and socially equitable (e.g. \u003csup\u003e28,29\u003c/sup\u003e). Furthermore, these MPAs are likely to help restore marine ecosystems and the recovery and long-term sustainability of fisheries \u003csup\u003e64,65\u003c/sup\u003e. Our pan-European assessment adds to growing evidence that the quality and level of protection within MPAs will be critical in determining whether quantitative targets such as protecting 30 % of European seas translate into meaningful, durable outcomes for both biodiversity, blue economies and people in a changing ocean.\u003c/p\u003e\n"},{"header":"Methods","content":"\u003cp\u003e\u003cem\u003eStudy region and MPAs\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThis study investigated protection levels at the pan-European scale, considering the EEZs of EU member states in addition to those of the Iceland, Norway (including Svalbard and Jan Mayen under Norwegian sovereignty), the Faroe Islands, and the United Kingdom (Figure 1). The resulting study region covered a total area of approximately 9,160,000 km\u003csup\u003e2\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003ePolygon shapefiles representing MPA boundaries were obtained for each constituent country: for EU member states from \u003csup\u003e25\u003c/sup\u003e, originally extracted from the European Environment Agency \u003csup\u003e66\u003c/sup\u003e (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 5,362); for the UK, polygons representing marine-associated Special Areas of Conservation (SAC), marine-associated Special Protection Areas (SPA), Nature Conservation Marine Protected Areas (NCMPA), and Marine Conservation Zones (MCZ) (including Highly Protected Marine Areas (HPMA)) (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 391) were obtained from various geospatial platforms of the UK\u0026apos;s Statutory Nature Conservation Bodies (SNCBs) \u003csup\u003e67\u003c/sup\u003e; Protected area shapefiles for Norway (\u003cem\u003en\u003c/em\u003e = 858) were obtained from \u003csup\u003e68\u003c/sup\u003e, while shapefiles for Iceland (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 68) and the Faroe Islands (\u003cem\u003en\u003c/em\u003e = 3) were obtained from the World Database on Protected Areas (WDPA) \u003csup\u003e69\u003c/sup\u003e and clipped to exclude areas on land.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eMPA protection level assignments\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe MPA Guide decision tree framework \u003csup\u003e6,59\u003c/sup\u003e was applied to classify each MPA or zone into one of four protection levels or as incompatible with conservation, based on regulations for seven activity types: mining, dredging and dumping, infrastructure, anchoring, fisheries, aquaculture and non-extractive uses \u003csup\u003e6\u003c/sup\u003e. The protection levels were: \u0026lsquo;Fully\u0026rsquo; protected, where all extractive or destructive activities are prohibited and other impacts are minimized; \u0026lsquo;Highly\u0026rsquo; protected, where only light extractive uses with low total impact are permitted; \u0026lsquo;Lightly\u0026rsquo; protected, where some biodiversity protection exists, but moderate extractive impacts are allowed; \u0026lsquo;Minimally\u0026rsquo; protected, allowing extensive extraction and high impact uses; Incompatible with conservation of nature, whereby activities deemed too impactful for biodiversity protection are allowed to occur. MPAs lacking sufficient regulatory information were labelled \u003cem\u003eUnclassified\u003c/em\u003e. \u0026lsquo;Fully\u0026rsquo; and \u0026lsquo;Highly\u0026rsquo; protected areas are expected to deliver the strongest biodiversity and ecosystem recovery benefits, with \u0026lsquo;Fully\u0026rsquo; protected MPAs offering the greatest gains, whereas \u0026lsquo;Lightly\u0026rsquo; and \u0026lsquo;Minimally\u0026rsquo; protected MPAs may yield limited or no species-specific benefits, with little effect on biodiversity at the ecosystem level, for example due to trophic cascades \u003csup\u003e59,62,70\u003c/sup\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFor EU member states, protection level classifications were already available from \u003csup\u003e25\u003c/sup\u003e.Aminian-Biquet et al., 2024 The EU MPA protection levels were available for two impact scenarios to account for situations where there was uncertainty regarding the scale and potential impacts of activities: the first assumed that, unless explicitly specified, the lowest possible impacts were occurring, while the second assumed the highest possible impacts were occurring \u003csup\u003e25\u003c/sup\u003e. For this study, assigned MPA protection levels assuming the lowest possible impacts were used, therefore in cases where there was uncertainty regarding the scale and potential impacts of activities the results represent an optimistic view of the levels of protection provided by the MPAs. Sufficient information was available for most UK and Norwegian MPAs to make informed decisions regarding the levels of impact for different activities, but where uncertainty did exist, the same approach was followed as for the EU MPAs \u003csup\u003e25,67,68\u003c/sup\u003e. There was insufficient information to assess the levels of protection provided by the Iceland and the Faroe Islands protected areas, therefore these were all categorised as \u0026lsquo;Unclassified\u0026rsquo;. Combined, the Icelandic and Faroese areas represent only 3,488 km\u003csup\u003e2\u0026nbsp;\u003c/sup\u003e(~0.3% of the total MPA area considered in this study) but the total marine area represented by their constituent national EEZs are approximately 11% of pan-European seas, therefore it was important to consider them in the analyses. All overlapping MPA polygons (i.e., resulting from multiple designations) with the same protection level were merged to remove areas of overlap and thereby avoid the double-counting of areas. In cases where overlapping MPA polygons represented different protection levels, the highest level of protection present was assigned to the overlap.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eSpecies distribution modelling\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe current distributions of marine species within national EEZs were obtained from species distribution models (SDMs) in \u003csup\u003e36\u003c/sup\u003e. These SDMs were developed using three different algorithms (Maxent, Random Forest, and XGBoost) under a point-process framework, and are also available for different time periods and Shared Socio-economic Pathway emissions scenarios.\u003c/p\u003e\n\u003cp\u003eFor the present analysis, we extracted species distributions for the present-day and for the \u0026lsquo;most-likely\u0026rsquo; SSP2-4.5 scenario for the year 2100. Where possible, the ensemble prediction was selected, but where the ensemble was not available the model with the highest performance across all methods (as measured by the Continuous Boyce Index, \u003csup\u003e71\u003c/sup\u003e) was selected. Species distribution layers were then clipped and masked to the area where the species was considered native. This was done by using the biogeographic realms from \u003csup\u003e72\u003c/sup\u003e; realms were overlaid with occurrence records, and those containing records were considered part of the native range. A buffer of 0.2 degrees was applied to the occurrence records to ensure that realms at the edges of species distributions were also included. We further restricted the distributions to the maximum recorded depth for each species, based on the occurrence data used to fit the models. The same process was applied to the future prediction layers.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003ePredictions were also uncertainty-discounted by subtracting the standard deviation across bootstraps from the average predicted probability of occurrence for each cell, using a fixed weight (0.5). Finally, we applied a threshold to convert probability of occurrence to presence or absence, using the model-derived threshold that maximized sensitivity (i.e. correctly identifying presences) and specificity (i.e. correctly identifying absences). This threshold has been shown to perform well even with presence-only models, such as this study, where no absence data is available and only background points are used \u003csup\u003e73,74\u003c/sup\u003e. As a final screening step, species with fewer than 50 predicted raster cells under present-day conditions were excluded from further analysis to avoid including taxa that are marginal or poorly predicted within the study area. Appropriate present-day distribution predictions were ultimately available for 9,794 species. Projected distributions under SSP2-4.5 were also extracted for these taxa, although slightly fewer were available (\u003cem\u003en\u0026nbsp;\u003c/em\u003e= 9,635) due to the processing steps above.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eBiotic group assignments\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eSpecies were classified into primary biotic groups based on their taxonomy and ecology. All photoautotrophs including macrophytes (e.g., seagrasses and macroalgae) and phytoplankton were assigned to the \u0026lsquo;Primary producers\u0026rsquo; group. Demersal and pelagic habitat assignments for fish were obtained based on their classification in FishBase (\u0026lsquo;Pelagic fish\u0026rsquo; were those species identified as \u0026lsquo;benthopelagic\u0026rsquo;, \u0026lsquo;bathypelagic\u0026rsquo;, \u0026lsquo;pelagic-oceanic\u0026rsquo;, \u0026lsquo;pelagic-neritic\u0026rsquo;, or \u0026lsquo;pelagic\u0026rsquo;; \u0026lsquo;Demersal fish\u0026rsquo; were those species identified as reef-associated\u0026rsquo;, \u0026lsquo;demersal\u0026rsquo;, or \u0026lsquo;bathydemersal\u0026rsquo;), using the \u0026lsquo;\u003cem\u003erfishbase\u003c/em\u003e\u0026rsquo; r package \u003csup\u003e75\u003c/sup\u003e. Invertebrates were split into either benthic or pelagic based on their taxonomy and known ecology in a top-down manner from class to species level: at each level, the relevant functional group classification from WoRMS was assigned to all taxa within that level (if available); assignments for any remaining taxa were then conducted at the next level (e.g. order), and so on until family level. For the few remaining taxa, assignments were then made at the species level. The invertebrates were then also screened to identify parasitic species, which were assigned to an \u0026lsquo;Others\u0026rsquo; group. The invertebrate order Limnomedusae (Class Hydrozoa) was also assigned to the group \u0026lsquo;Others\u0026rsquo; due to their combined benthic and pelagic lifecycle. Taxa deemed not to be strictly marine (birds and marine-associated terrestrial invertebrates), and reptiles (all turtles) were also assigned to this group. Due to the mixed membership of this \u0026lsquo;Others\u0026rsquo; group, all constituent species (\u003cem\u003en\u003c/em\u003e = 1,028) were excluded from further analyses.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTo explore how well MPA classes cover the distributions of key taxa, species were also separately grouped based on their conservation status and whether they are important habitat-formers. Species were classified as either threatened or non-threatened, with the former made up of those listed on the IUCN Red List as Vulnerable, Endangered or Critically endangered (see further details in \u003csup\u003e76\u003c/sup\u003e). The habitat forming species were those categorized into one of nine distinct biogenic habitat groups: bryozoan reefs; cold-water coral reefs; coralligenous platforms; macroalgae forests; deep-sea sponge grounds; mollusc reefs; polychaete reefs; seagrasses; and shallow-water sponge reefs \u003csup\u003e77\u003c/sup\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eAnalyses\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eOverlap between MPA protection levels and species\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eAll analyses were conducted in R version 4.5.0 \u003csup\u003e78\u003c/sup\u003e. Rasters were handled using the \u003cem\u003eterra\u0026nbsp;\u003c/em\u003epackage \u003csup\u003e79\u003c/sup\u003e. For each species, the total area of overlap between cells with predicted presences and each MPA protection level was determined using the \u003cem\u003eexactextractr\u003c/em\u003e package \u003csup\u003e80\u003c/sup\u003e. This approach facilitates the calculation of zonal statistics for rasters covered by polygon areas, including areas of partial overlap \u003csup\u003e80\u003c/sup\u003e. The proportion of each species\u0026rsquo; range (i.e., the summed area of cells with predicted presences) covered by each MPA protection level was then calculated. This process was conducted for the predicted spatial distributions under both present-day and future (SSP2-4.5) environmental conditions.\u003c/p\u003e\n\u003cp\u003eThe distributions of MPA protection level coverage across all species and for each biotic group (including threatened and habitat-forming taxa) under present-day environmental conditions were visualised using violin plots and boxplots, and described using summary statistics (medians, 95\u003csup\u003eth\u003c/sup\u003e percentiles and minimum/maximum). When reporting results in text, the two highest protection levels (\u0026lsquo;Fully\u0026rsquo; and \u0026lsquo;Highly\u0026rsquo;) are referred to as a single combined level \u003cem\u003estrong protection\u003c/em\u003e, and the remaining levels (excluding \u0026lsquo;Unclassified\u0026rsquo;) were combined into a \u003cem\u003eweak protection\u003c/em\u003e level, following \u003csup\u003e25\u003c/sup\u003e. This was done to clearly differentiate the protection levels which are the most likely to have positive outcomes for biodiversity \u003csup\u003e6\u003c/sup\u003e. Protection levels are reported individually in figures except for \u0026lsquo;Fully\u0026rsquo; and \u0026lsquo;Highly\u0026rsquo; which represent low overall areas (~1% of all pan-European MPAs) and were therefore grouped into \u0026ldquo;Fully / Highly\u0026rdquo; for visual clarity. Summary statistics in tables are reported separately for each MPA protection level.\u003c/p\u003e\n\u003cp\u003eTo explore the climate readiness of the pan-European MPAs, the overlap of future species distributions with MPAs was calculated using the same approach as that described for present-day distributions. The relative changes in the areas of overlap for each species and MPA protection level between current predicted distributions and projected future distributions were also calculated, by dividing the future projected areas of coverage by the current day predicted areas of coverage.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAs a simple indication of how climate change may impact species both directly through expansions or contractions in total range area and indirectly through shifts in distribution and associated coverage by MPAs, species were split into risk categories based on their relative changes in total range area (e.g., see \u003csup\u003e32\u003c/sup\u003e): species projected to experience distribution contractions of \u0026gt;50% were considered \u0026lsquo;High climate-risk\u0026rsquo;; species projected to lose between 10% and 50% of their distribution were considered \u0026lsquo;Medium climate-risk\u0026rsquo;; those projected to lose \u0026lt;10% of their distribution or experience net gains were considered \u0026lsquo;Low climate-risk\u0026rsquo;. The distribution of relative changes in areas of overlap with MPA protection levels was summarised by biotic group for each risk class separately.\u0026nbsp;\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData and code will be made available in an open-access repository (e.g. Zenodo) upon acceptance of this manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eF.S. and P.A.E-K were funded through the Newcastle University One Planet NUAcT Fellowship. F.S., P.A., and D.S. acknowledge support from Oceana for work related to Marine Protected Areas in a Changing Climate. A.M.A., S.C.P., J.A., and M.J.C. acknowledge support from the MPA Europe (HORIZON-CL6-2023-BIODIV-01-12, Grant Agreement No. 101059988), co-funded by the European Union\u0026rsquo;s Horizon Europe Programme. B.H.C., I.S., J.M.A., E.M., and A.R.K. acknowledge support from the MARHAB project (HORIZON-CL6-2023-BIODIV-01-4, Grant Agreement No. 101135307), funded by the European Union\u0026rsquo;s Horizon Europe Programme. B.H.C., I.S., and J.A. also received support from Portuguese national funds from FCT \u0026ndash; Foundation for Science and Technology through contracts UID/04326/2025 (DOI: https://doi.org/10.54499/UID/04326/2025), UID/PRR/04326/2025 (DOI: https://doi.org/10.54499/UID/PRR/04326/2025), and LA/P/0101/2020 (DOI: https://doi.org/10.54499/LA/P/0101/2020) to CCMAR. B.H.C. also acknowledges support from the MarPlus2024 (Algarve 2030, ALGARVE-FSE+-01177700) project. The authors also thank Beth Pike, Jenna Sullivan-Stack, Kirsten Grorud-Colvert, and Joachim Claudet for valuable discussions.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCRediT statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePatrick A. Eskuche-Keith: Conceptualisation, Data Curation, Methodology, Investigation, Formal analysis, Visualisation, Writing - original draft, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eFabrice Stephenson: Conceptualisation, Supervision, Funding acquisition, Methodology, Investigation, Writing - original draft, Writing - Review \u0026amp; Editing\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eMark J. Costello: Conceptualisation, Supervision, Funding acquisition, Methodology, Writing - original draft, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003ePippa J. Moore: Conceptualisation, Funding acquisition, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eB\u0026aacute;rbara Horta e Costa: Conceptualisation, Funding acquisition, Supervision, Data Curation, Methodology, Writing - original draft, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eAnna M. Addamo: Conceptualisation, Methodology, Data Curation, Writing - original draft, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eSilas C. Principe: Conceptualisation, Methodology, Data Curation, Formal analysis, Writing - original draft, Investigation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003ePeter M. Almond: Writing - original draft, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eJorge Assis: Conceptualisation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eIn\u0026ecirc;s Sousa: Investigation, Data Curation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eEven Moland: Investigation, Data Curation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eAlf R. Kleiven: Investigation, Data Curation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eJohanna M. Aarflot: Investigation, Data Curation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eElsa Sim: Investigation, Data Curation, Writing - Review \u0026amp; Editing\u003c/p\u003e\n\u003cp\u003eDaniel Skerritt: Conceptualisation, Writing - Review \u0026amp; Editing\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eJaureguiberry, P.\u003cem\u003e et al.\u003c/em\u003e The direct drivers of recent global anthropogenic biodiversity loss. \u003cem\u003eScience Advances\u003c/em\u003e \u003cstrong\u003e8\u003c/strong\u003e (2022). https://doi.org/10.1126/sciadv.abm9982\u003c/li\u003e\n\u003cli\u003eHalpern, B. S., Frazier, M., O\u0026rsquo;Hara, C. C., Vargas-Fonseca, O. A. \u0026amp; Lombard, A. T. Cumulative impacts to global marine ecosystems projected to more than double by mid-century. \u003cem\u003eScience\u003c/em\u003e \u003cstrong\u003e389\u003c/strong\u003e, 1216-1219 (2025). https://doi.org/10.1126/science.adv2906\u003c/li\u003e\n\u003cli\u003eChaudhary, C., Richardson, A. J., Schoeman, D. S. \u0026amp; Costello, M. J. Global warming is causing a more pronounced dip in marine species richness around the equator. \u003cem\u003eProceedings of the National Academy of Sciences\u003c/em\u003e \u003cstrong\u003e118\u003c/strong\u003e, e2015094118 (2021). https://doi.org/10.1073/pnas.2015094118\u003c/li\u003e\n\u003cli\u003eHe, Q. \u0026amp; Silliman, B. R. Climate Change, Human Impacts, and Coastal Ecosystems in the Anthropocene. \u003cem\u003eCurrent Biology\u003c/em\u003e \u003cstrong\u003e29\u003c/strong\u003e, R1021-R1035 (2019). https://doi.org/10.1016/j.cub.2019.08.042\u003c/li\u003e\n\u003cli\u003eCostello, M. J. Long live Marine Reserves: A review of experiences and benefits. \u003cem\u003eBiological Conservation\u003c/em\u003e \u003cstrong\u003e176\u003c/strong\u003e, 289-296 (2014). https://doi.org/10.1016/j.biocon.2014.04.023\u003c/li\u003e\n\u003cli\u003eGrorud-Colvert, K.\u003cem\u003e et al.\u003c/em\u003e The MPA Guide: A framework to achieve global goals for the ocean. \u003cem\u003eScience\u003c/em\u003e \u003cstrong\u003e373\u003c/strong\u003e (2021). https://doi.org/10.1126/science.abf0861\u003c/li\u003e\n\u003cli\u003eGon\u0026ccedil;alves, E. J. Marine Protected Areas as Tools for Ocean Sustainability, 131-141, Springer International Publishing (2023).\u003c/li\u003e\n\u003cli\u003eWhite, J. W.\u003cem\u003e et al.\u003c/em\u003e Measurements, mechanisms, and management recommendations for how marine protected areas can provide climate resilience. \u003cem\u003eMarine Policy\u003c/em\u003e \u003cstrong\u003e171\u003c/strong\u003e, 106419 (2025). https://doi.org/10.1016/j.marpol.2024.106419\u003c/li\u003e\n\u003cli\u003eStephens, T. The Kunming\u0026ndash;Montreal Global Biodiversity Framework. \u003cem\u003eInternational Legal Materials\u003c/em\u003e \u003cstrong\u003e62\u003c/strong\u003e, 868-887 (2023). https://doi.org/10.1017/ilm.2023.16\u003c/li\u003e\n\u003cli\u003eWenzel, L.\u003cem\u003e et al.\u003c/em\u003e Establishing Marine Protected Areas in a Changing Climate. IUCN WCPA Technical Report Series No. 9., (2025).\u003c/li\u003e\n\u003cli\u003eKlein, C. J.\u003cem\u003e et al.\u003c/em\u003e Shortfalls in the global protected area network at representing marine biodiversity. \u003cem\u003eScientific Reports 2015 5:1\u003c/em\u003e \u003cstrong\u003e5\u003c/strong\u003e (2015). https://doi.org/10.1038/srep17539\u003c/li\u003e\n\u003cli\u003eAppeltans, W.\u003cem\u003e et al.\u003c/em\u003e Biodiversity knowledge and threats on marine life: assessing no-take zones as a refuge for marine species. (IOC-UNESCO, 2024).\u003c/li\u003e\n\u003cli\u003eZhao, Q. \u0026amp; Costello, M. J. Ecologically representative Marine Protected Area planning can think globally and act locally. \u003cem\u003eTrends in Ecology \u0026amp; Evolution\u003c/em\u003e \u003cstrong\u003e40\u003c/strong\u003e, 772-781 (2025). https://doi.org/10.1016/j.tree.2025.05.007\u003c/li\u003e\n\u003cli\u003eEdgar, G. J.\u003cem\u003e et al.\u003c/em\u003e Global conservation outcomes depend on marine protected areas with five key features. \u003cem\u003eNature\u003c/em\u003e \u003cstrong\u003e506\u003c/strong\u003e, 216-220 (2014). https://doi.org/10.1038/nature13022\u003c/li\u003e\n\u003cli\u003eDureuil, M., Boerder, K., Burnett, K. A., Froese, R. \u0026amp; Worm, B. Elevated trawling inside protected areas undermines conservation outcomes in a global fishing hot spot. \u003cem\u003eScience\u003c/em\u003e \u003cstrong\u003e362\u003c/strong\u003e, 1403-1407 (2018). https://doi.org/10.1126/science.aau0561\u003c/li\u003e\n\u003cli\u003eCostello, M. J. \u0026amp; Ballantine, B. Biodiversity conservation should focus on no-take Marine Reserves: 94% of Marine Protected Areas allow fishing. \u003cem\u003eTrends in Ecology \u0026amp; Evolution\u003c/em\u003e \u003cstrong\u003e30\u003c/strong\u003e, 507-509 (2015). https://doi.org/10.1016/j.tree.2015.06.011\u003c/li\u003e\n\u003cli\u003eBarnes, M. D., Glew, L., Wyborn, C. \u0026amp; Craigie, I. D. Prevent perverse outcomes from global protected area policy. \u003cem\u003eNature Ecology \u0026amp; Evolution\u003c/em\u003e \u003cstrong\u003e2\u003c/strong\u003e, 759-762 (2018). https://doi.org/10.1038/s41559-018-0501-y\u003c/li\u003e\n\u003cli\u003ePike, E. P.\u003cem\u003e et al.\u003c/em\u003e Ocean protection quality is lagging behind quantity: Applying a scientific framework to assess real marine protected area progress against the 30 by 30 target. \u003cem\u003eConservation Letters\u003c/em\u003e \u003cstrong\u003e17\u003c/strong\u003e (2024). https://doi.org/10.1111/conl.13020\u003c/li\u003e\n\u003cli\u003eCouncil of Europe. Convention on the Conservation of European Wildlife and Natural Habitats. CETS 104, Bern, (1979).\u003c/li\u003e\n\u003cli\u003eEuropean Council. Directive 2009/147/EC of the European Parliament and of the Council on the Conservation of Wild Birds (2009).\u003c/li\u003e\n\u003cli\u003eCentre for International Law. Convention on the Protection of the Black Sea Against Pollution (Bucharest Convention), Bucharest (1992).\u003c/li\u003e\n\u003cli\u003eRaftopoulos, E. The Barcelona Convention System for the Protection of the Mediterranean Sea against Pollution: An international trust at work. Int\u0026apos;l J. Estuarine \u0026amp; Coastal L., 7, p.27. \u003cem\u003eInternational Journal of Estuarine and Coastal Law\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e, 27-42 (1992). \u003c/li\u003e\n\u003cli\u003eTromp, D. \u0026amp; Wieriks, K. The OSPAR Convention: 25 years of North Sea protection. \u003cem\u003eMarine Pollution Bulletin\u003c/em\u003e \u003cstrong\u003e29\u003c/strong\u003e, 622-626 (1994). https://doi.org/10.1016/0025-326x(94)90698-x\u003c/li\u003e\n\u003cli\u003eFitzmaurice, M. The Helsinki Conventions 1974 and 1992. \u003cem\u003eThe International Journal of Marine and Coastal Law\u003c/em\u003e \u003cstrong\u003e13\u003c/strong\u003e, 379-394 (1998). \u003c/li\u003e\n\u003cli\u003eAminian-Biquet, J.\u003cem\u003e et al.\u003c/em\u003e Over 80% of the European Union\u0026rsquo;s marine protected area only marginally regulates human activities. \u003cem\u003eOne Earth\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e, 1614-1629 (2024). https://doi.org/10.1016/j.oneear.2024.07.010\u003c/li\u003e\n\u003cli\u003eEuropean Commission. EU Biodiversity Strategy for 2030: Bringing nature back into our lives. Brussels. Available at: https://eur-lex.europa.eu/legal-content/EN/TXT/?uri=CELEX:52020DC0380 (2020).\u003c/li\u003e\n\u003cli\u003eEuropean Commission. Criteria and guidance for protected areas designations. Brussels. Available at: https://environment.ec.europa.eu/publications/criteria-and-guidance-protected-areas-designations-staff-working-document_en (2022).\u003c/li\u003e\n\u003cli\u003eHorta e Costa, B., Stephenson, F. \u0026amp; Claudet, J. European Union\u0026rsquo;s strict conservation targets should guide global marine policy. \u003cem\u003eNature\u003c/em\u003e \u003cstrong\u003e642\u003c/strong\u003e, 38-38 (2025). https://doi.org/10.1038/d41586-025-01731-7\u003c/li\u003e\n\u003cli\u003eStephenson, F.\u003cem\u003e et al.\u003c/em\u003e Quality of marine protected areas is critical to achieving global biodiversity targets. \u003cem\u003enpj Ocean Sustainability\u003c/em\u003e \u003cstrong\u003e4\u003c/strong\u003e (2025). https://doi.org/10.1038/s44183-025-00169-8\u003c/li\u003e\n\u003cli\u003eAmengual, J. \u0026amp; Alvarez-Berastegui, D. A critical evaluation of the Aichi Biodiversity Target 11 and the Mediterranean MPA network, two years ahead of its deadline. \u003cem\u003eBiological Conservation\u003c/em\u003e \u003cstrong\u003e225\u003c/strong\u003e, 187-196 (2018). https://doi.org/10.1016/j.biocon.2018.06.032\u003c/li\u003e\n\u003cli\u003eEEA. State of Europe\u0026rsquo;s Seas, Technical report No. 2., Copenhagen (2015).\u003c/li\u003e\n\u003cli\u003eStephenson, F.\u003cem\u003e et al.\u003c/em\u003e Implications for the conservation of deep-water corals in the face of multiple stressors: A case study from the New Zealand region. \u003cem\u003eJournal of Environmental Management\u003c/em\u003e \u003cstrong\u003e346\u003c/strong\u003e, 118938 (2023). https://doi.org/10.1016/j.jenvman.2023.118938\u003c/li\u003e\n\u003cli\u003eZelli, E.\u003cem\u003e et al.\u003c/em\u003e Identifying climate refugia for vulnerable marine ecosystem indicator taxa under future climate change scenarios. \u003cem\u003eJournal of Environmental Management\u003c/em\u003e \u003cstrong\u003e373\u003c/strong\u003e, 122635 (2025). https://doi.org/10.1016/j.jenvman.2024.122635\u003c/li\u003e\n\u003cli\u003eOBIS. Ocean Biodiversity Information System. Intergovernmental Oceanographic Commission of UNESCO (2026).\u003c/li\u003e\n\u003cli\u003eAssis, J.\u003cem\u003e et al.\u003c/em\u003e Bio‐ORACLE v3.0. Pushing marine data layers to the CMIP6 Earth System Models of climate change research. \u003cem\u003eGlobal Ecology and Biogeography\u003c/em\u003e \u003cstrong\u003e33\u003c/strong\u003e (2024). https://doi.org/10.1111/geb.13813\u003c/li\u003e\n\u003cli\u003ePrincipe, S. C.\u003cem\u003e et al.\u003c/em\u003e Mapping marine species distributions to inform the design of protected areas in Europe [Manuscript submitted for publication]. \u003cem\u003eScientific Data\u003c/em\u003e (2026). \u003c/li\u003e\n\u003cli\u003eAssis, J., Fragkopoulou, E., Gouv\u0026ecirc;a, L., Ara\u0026uacute;jo, M. B. \u0026amp; Serr\u0026atilde;o, E. A. Kelp forest diversity under projected end‐of‐century climate change. \u003cem\u003eDiversity and Distributions\u003c/em\u003e \u003cstrong\u003e30\u003c/strong\u003e (2024). https://doi.org/10.1111/ddi.13837\u003c/li\u003e\n\u003cli\u003eGidden, M. J.\u003cem\u003e et al.\u003c/em\u003e Global emissions pathways under different socioeconomic scenarios for use in CMIP6: a dataset of harmonized emissions trajectories through the end of the century. \u003cem\u003eGeoscientific Model Development\u003c/em\u003e \u003cstrong\u003e12\u003c/strong\u003e, 1443-1475 (2019). https://doi.org/10.5194/gmd-12-1443-2019\u003c/li\u003e\n\u003cli\u003eDavies, T. E., Maxwell, S. M., Kaschner, K., Garilao, C. \u0026amp; Ban, N. C. Large marine protected areas represent biodiversity now and under climate change. \u003cem\u003eScientific Reports\u003c/em\u003e \u003cstrong\u003e7\u003c/strong\u003e (2017). https://doi.org/10.1038/s41598-017-08758-5\u003c/li\u003e\n\u003cli\u003eGiakoumi, S.\u003cem\u003e et al.\u003c/em\u003e Revisiting \u0026ldquo;Success\u0026rdquo; and \u0026ldquo;Failure\u0026rdquo; of Marine Protected Areas: A Conservation Scientist Perspective. \u003cem\u003eFrontiers in Marine Science\u003c/em\u003e \u003cstrong\u003e5\u003c/strong\u003e (2018). https://doi.org/10.3389/fmars.2018.00223\u003c/li\u003e\n\u003cli\u003eBlowes, S. A.\u003cem\u003e et al.\u003c/em\u003e Mediterranean marine protected areas have higher biodiversity via increased evenness, not abundance. \u003cem\u003eJournal of Applied Ecology\u003c/em\u003e \u003cstrong\u003e57\u003c/strong\u003e, 578-589 (2020). https://doi.org/10.1111/1365-2664.13549\u003c/li\u003e\n\u003cli\u003eSullivan-Stack, J.\u003cem\u003e et al.\u003c/em\u003e Assessments of expected MPA outcomes can inform and improve biodiversity conservation: Case studies using The MPA Guide. \u003cem\u003eMarine Policy\u003c/em\u003e \u003cstrong\u003e170\u003c/strong\u003e, 106364 (2024). https://doi.org/10.1016/j.marpol.2024.106364\u003c/li\u003e\n\u003cli\u003eMoilanen, A., Kujala, H. \u0026amp; Leathwick, J. R. The Zonation Framework and Software for Conservation Prioritization. 196-210, Oxford University Press, Oxford (2009).\u003c/li\u003e\n\u003cli\u003eZhao, Q., Huang, H. \u0026amp; Costello, M. J. Systematic planning shows more than half of the most species-rich ocean region is needed to include all species in representative protected areas. \u003cem\u003eGlobal Ecology and Conservation\u003c/em\u003e \u003cstrong\u003e53\u003c/strong\u003e, e03036 (2024). https://doi.org/10.1016/j.gecco.2024.e03036\u003c/li\u003e\n\u003cli\u003eLauer, D. \u0026amp; Reaka, M. Depth distributions of benthic and pelagic species highlight the potential of mesophotic and deep habitats to serve as marine refugia. \u003cem\u003eMarine Ecology Progress Series\u003c/em\u003e \u003cstrong\u003e700\u003c/strong\u003e, 39-52 (2022). https://doi.org/10.3354/meps14180\u003c/li\u003e\n\u003cli\u003eMarrocco, V.\u003cem\u003e et al.\u003c/em\u003e Behavioural Constraints to Home Range Allometries in Aquatic Organisms. \u003cem\u003eEcology and Evolution\u003c/em\u003e \u003cstrong\u003e15\u003c/strong\u003e (2025). https://doi.org/10.1002/ece3.71886\u003c/li\u003e\n\u003cli\u003eM\u0026ouml;llmann, C., M\u0026uuml;ller-Karulis, B., Kornilovs, G. \u0026amp; St John, M. A. Effects of climate and overfishing on zooplankton dynamics and ecosystem structure: regime shifts, trophic cascade, and feedback loops in a simple ecosystem. \u003cem\u003eICES Journal of Marine Science\u003c/em\u003e \u003cstrong\u003e65\u003c/strong\u003e, 302-310 (2008). https://doi.org/10.1093/icesjms/fsm197\u003c/li\u003e\n\u003cli\u003eRoberts, K. E., Smith, B. J., Burkholder, D. \u0026amp; Hart, K. M. Evaluating the use of marine protected areas by endangered species: A habitat selection approach. \u003cem\u003eEcological Solutions and Evidence\u003c/em\u003e \u003cstrong\u003e2\u003c/strong\u003e (2021). https://doi.org/10.1002/2688-8319.12035\u003c/li\u003e\n\u003cli\u003eGilmour, M. E.\u003cem\u003e et al.\u003c/em\u003e Evaluation of MPA designs that protect highly mobile megafauna now and under climate change scenarios. \u003cem\u003eGlobal Ecology and Conservation\u003c/em\u003e \u003cstrong\u003e35\u003c/strong\u003e, e02070 (2022). https://doi.org/10.1016/j.gecco.2022.e02070\u003c/li\u003e\n\u003cli\u003ePredragovic, M.\u003cem\u003e et al.\u003c/em\u003e Up to 80% of threatened and commercial species across European marine protected areas face novel climates under high emission scenario. \u003cem\u003enpj Ocean Sustainability\u003c/em\u003e \u003cstrong\u003e3\u003c/strong\u003e (2024). https://doi.org/10.1038/s44183-024-00068-4\u003c/li\u003e\n\u003cli\u003ePoloczanska, E. S.\u003cem\u003e et al.\u003c/em\u003e Global imprint of climate change on marine life. \u003cem\u003eNature Climate Change\u003c/em\u003e \u003cstrong\u003e3\u003c/strong\u003e, 919-925 (2013). https://doi.org/10.1038/nclimate1958\u003c/li\u003e\n\u003cli\u003eBurrows, M. T.\u003cem\u003e et al.\u003c/em\u003e Ocean community warming responses explained by thermal affinities and temperature gradients. \u003cem\u003eNature Climate Change\u003c/em\u003e \u003cstrong\u003e9\u003c/strong\u003e, 959-963 (2019). https://doi.org/10.1038/s41558-019-0631-5\u003c/li\u003e\n\u003cli\u003eGord\u0026oacute;-Vilaseca, C., Stephenson, F., Coll, M., Lavin, C. \u0026amp; Costello, M. J. Three decades of increasing fish biodiversity across the northeast Atlantic and the Arctic Ocean. \u003cem\u003eProceedings of the National Academy of Sciences\u003c/em\u003e \u003cstrong\u003e120\u003c/strong\u003e (2023). https://doi.org/10.1073/pnas.2120869120\u003c/li\u003e\n\u003cli\u003eGord\u0026oacute;-Vilaseca, C.\u003cem\u003e et al.\u003c/em\u003e Future trends of marine fish biomass distributions from the North Sea to the Barents Sea. \u003cem\u003eNature Communications 2024 15:1\u003c/em\u003e \u003cstrong\u003e15\u003c/strong\u003e (2024). https://doi.org/10.1038/s41467-024-49911-9\u003c/li\u003e\n\u003cli\u003eBartlett, B. S., Erisman, B. \u0026amp; Asch, R. G. Current Marine Protected Areas Conserve Fish Spawning Aggregations Under Climate Change due to Habitat Refugia. \u003cem\u003eGlobal Change Biology\u003c/em\u003e \u003cstrong\u003e31\u003c/strong\u003e (2025). https://doi.org/10.1111/gcb.70433\u003c/li\u003e\n\u003cli\u003eMalanoski, C. M., Farnsworth, A., Lunt, D. J., Valdes, P. J. \u0026amp; Saupe, E. E. Climate change is an important predictor of extinction risk on macroevolutionary timescales. \u003cem\u003eScience\u003c/em\u003e \u003cstrong\u003e383\u003c/strong\u003e, 1130-1134 (2024). https://doi.org/10.1126/science.adj5763\u003c/li\u003e\n\u003cli\u003eManes, S.\u003cem\u003e et al.\u003c/em\u003e Endemism increases species\u0026apos; climate change risk in areas of global biodiversity importance. \u003cem\u003eBiological Conservation\u003c/em\u003e \u003cstrong\u003e257\u003c/strong\u003e, 109070 (2021). https://doi.org/10.1016/j.biocon.2021.109070\u003c/li\u003e\n\u003cli\u003eMazaris, A. D.\u003cem\u003e et al.\u003c/em\u003e Threats to marine biodiversity in European protected areas. \u003cem\u003eScience of The Total Environment\u003c/em\u003e \u003cstrong\u003e677\u003c/strong\u003e, 418-426 (2019). https://doi.org/10.1016/j.scitotenv.2019.04.333\u003c/li\u003e\n\u003cli\u003eOregon State University, IUCN World Commission on Protected Areas - Marine, Marine Conservation Institute, National Geographic Pristine Seas \u0026amp; UN Environment Programme World Conservation Monitoring Centre. The MPA Guide User Manual, version 1., https://mpa-guide.protectedplanet.net (2023).\u003c/li\u003e\n\u003cli\u003eHermoso, V.\u003cem\u003e et al.\u003c/em\u003e The EU Biodiversity Strategy for 2030: Opportunities and challenges on the path towards biodiversity recovery. \u003cem\u003eEnvironmental Science \u0026amp; Policy\u003c/em\u003e \u003cstrong\u003e127\u003c/strong\u003e, 263-271 (2022). https://doi.org/10.1016/j.envsci.2021.10.028\u003c/li\u003e\n\u003cli\u003eGill, D. A.\u003cem\u003e et al.\u003c/em\u003e Capacity shortfalls hinder the performance of marine protected areas globally. \u003cem\u003eNature\u003c/em\u003e \u003cstrong\u003e543\u003c/strong\u003e, 665-669 (2017). https://doi.org/10.1038/nature21708\u003c/li\u003e\n\u003cli\u003eHorta e Costa, B.\u003cem\u003e et al.\u003c/em\u003e Marine protected areas stage of establishment and level of protection are good predictors of their conservation outcomes. \u003cem\u003eCell Reports Sustainability\u003c/em\u003e \u003cstrong\u003e2\u003c/strong\u003e, 100345 (2025). https://doi.org/10.1016/j.crsus.2025.100345\u003c/li\u003e\n\u003cli\u003eZucchetta, M.\u003cem\u003e et al.\u003c/em\u003e Can the Effects of Anthropogenic Pressures and Environmental Variability on Nekton Fauna Be Detected in Fishery Data? Insights from the Monitoring of the Artisanal Fishery Within the Venice Lagoon. \u003cem\u003eEstuaries and Coasts\u003c/em\u003e \u003cstrong\u003e39\u003c/strong\u003e, 1164-1182 (2016). https://doi.org/10.1007/s12237-015-0064-y\u003c/li\u003e\n\u003cli\u003eCostello, M. J. Evidence of economic benefits from marine protected areas. \u003cem\u003eScientia Marina\u003c/em\u003e \u003cstrong\u003e88\u003c/strong\u003e, e080 (2024). https://doi.org/10.3989/scimar.05417.080\u003c/li\u003e\n\u003cli\u003eCostello, M. J. Fully protected Marine Protected Areas do not displace fisheries. \u003cem\u003eProceedings of the National Academy of Sciences\u003c/em\u003e \u003cstrong\u003e121\u003c/strong\u003e (2024). https://doi.org/10.1073/pnas.2412543121\u003c/li\u003e\n\u003cli\u003eEuropean Environment Agency. Marine Protected Areas (MPA) in EEA marine assessment areas, Ver. 2021. In: AGENCY, E. E. (ed.). DOI: eea_v_3035_100_k_mpa-in-marine-assessment-area_p_2021_v01_r00 (2023).\u003c/li\u003e\n\u003cli\u003eSim, E. Regulations of Human Activities \u0026amp; Protection Levels in Marine Protected Areas of the United Kingdom. [dataset], Figshare (2025). https://doi.org/10.6084/m9.figshare.29901479.v5. \u003c/li\u003e\n\u003cli\u003eHorta e Costa, B., Sousa, I., Kleiven, A. R., Aarflot, J. M. \u0026amp; Moland, E. Data on Marine Protection Levels in Norwegian MPAs: A National Application of The MPA Guide. \u003cem\u003eData in Brief\u003c/em\u003e (submitted). \u003c/li\u003e\n\u003cli\u003eUNEP-WCMC and IUCN. Protected Planet: The World Database on Protected Areas (WDPA) (2026). https://doi.org/10.34892/6fwd-af11. \u003c/li\u003e\n\u003cli\u003eCostello, M. J., Gord\u0026oacute;-Vilaseca, C. \u0026amp; Coll, M. Trophic Cascades and Marine Reserves: Dual Indicators of Fishery and Climate Change Disruption in Pelagic and Benthic Ecosystems. \u003cem\u003eIn: Imperiled: The Encyclopedia of Conservation\u003c/em\u003e. 903-911, Elsevier (2022).\u003c/li\u003e\n\u003cli\u003eHirzel, A. H., Le Lay, G., Helfer, V., Randin, C. \u0026amp; Guisan, A. Evaluating the ability of habitat suitability models to predict species presences. \u003cem\u003eEcological Modelling\u003c/em\u003e \u003cstrong\u003e199\u003c/strong\u003e, 142-152 (2006). https://doi.org/10.1016/j.ecolmodel.2006.05.017\u003c/li\u003e\n\u003cli\u003eCostello, M. J.\u003cem\u003e et al.\u003c/em\u003e Marine biogeographic realms and species endemicity. \u003cem\u003eNature Communications\u003c/em\u003e \u003cstrong\u003e8\u003c/strong\u003e (2017). https://doi.org/10.1038/s41467-017-01121-2\u003c/li\u003e\n\u003cli\u003eLiu, C., Newell, G. \u0026amp; White, M. On the selection of thresholds for predicting species occurrence with presence‐only data. \u003cem\u003eEcology and Evolution\u003c/em\u003e \u003cstrong\u003e6\u003c/strong\u003e, 337-348 (2016). https://doi.org/10.1002/ece3.1878\u003c/li\u003e\n\u003cli\u003eLiu, C., White, M. \u0026amp; Newell, G. Selecting thresholds for the prediction of species occurrence with presence‐only data. \u003cem\u003eJournal of Biogeography\u003c/em\u003e \u003cstrong\u003e40\u003c/strong\u003e, 778-789 (2013). https://doi.org/10.1111/jbi.12058\u003c/li\u003e\n\u003cli\u003eBoettiger, C., Lang, D. T. \u0026amp; Wainwright, P. C. rfishbase: exploring, manipulating and visualizing FishBase data from R. \u003cem\u003eJournal of Fish Biology\u003c/em\u003e \u003cstrong\u003e81\u003c/strong\u003e, 2030-2039 (2012). https://doi.org/10.1111/j.1095-8649.2012.03464.x\u003c/li\u003e\n\u003cli\u003ePrincipe, S.\u003cem\u003e et al.\u003c/em\u003e MPA Europe Deliverable 3.3. Database of species and habitats conservation status. Zenodo (2024). https://doi.org/10.5281/zenodo.11075390\u003c/li\u003e\n\u003cli\u003ePrincipe, S.\u003cem\u003e et al.\u003c/em\u003e Publish maps and models of biogenic habitat distribution in Europe on EMODnet. Zenodo (2023). https://doi.org/10.5281/zenodo.10422129\u003c/li\u003e\n\u003cli\u003eR Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, Vienna, Austria, 2025). https://www.R-project.org/.\u003c/li\u003e\n\u003cli\u003eHijmans. terra: Spatial Data Analysis. R package version 1.8-93. https://cran.r-project.org/web/packages/terra/index.html (2026).\u003c/li\u003e\n\u003cli\u003eBaston. exactextractr: Fast Extraction from Raster Datasets using Polygons. R package version 0.10.0 https://CRAN.R-project.org/package=exactextractr (2023).\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"nature-portfolio","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"","title":"Nature Portfolio","twitterHandle":"","acdcEnabled":false,"dfaEnabled":false,"editorialSystem":"ejp","reportingPortfolio":"","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Species distribution modelling, marine spatial planning, area-based conservation, range shifts, management effectiveness, marine policy","lastPublishedDoi":"10.21203/rs.3.rs-8894972/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8894972/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Marine Protected Area (MPA) effectiveness depends on both coverage of species distributions and protection levels. It remains unclear whether Europe’s expanding MPA network adequately represents biodiversity now and as species redistribute under climate change. We provide the first pan-European assessment integrating MPA Guide protection levels with modelled present and future distributions for over 9,000 marine species. MPAs currently cover ~11 % of pan-European seas but encompass ~25% of species’ ranges, indicating preferential placement in biodiversity-rich areas. Representation remains stable under end-century projections (SSP2-4.5), and species at high climate risk generally retain or increase MPA coverage, suggesting protection of climatic refugia. However, \u003c 1 % of species’ distributions fall within areas restricting harmful practices such as fishing. Europe’s MPA network is well placed for biodiversity conservation and climate resilience but weakly protected, limiting its conservation impact and highlighting the need to strengthen management and strategically expand effectively protected and climate-resilient areas.","manuscriptTitle":"European Marine Protected Areas are well-placed and climate-resilient, but weakly protected","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-09 08:13:08","doi":"10.21203/rs.3.rs-8894972/v1","editorialEvents":[],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"nature-communications","isNatureJournal":true,"hasQc":false,"allowDirectSubmit":false,"externalIdentity":"NCOMMS","sideBox":"Learn more about [Nature Communications](http://www.nature.com/ncomms/)","snPcode":"","submissionUrl":"https://mts-ncomms.nature.com/","title":"Nature Communications","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"ejp","reportingPortfolio":"Nature Communications","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"2f3d1261-a0de-4666-ae6f-6464dd6eaa5e","owner":[],"postedDate":"April 9th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[{"id":64471615,"name":"Biological sciences/Ecology/Conservation biology"},{"id":64471616,"name":"Biological sciences/Ecology/Biogeography"},{"id":64471617,"name":"Biological sciences/Ecology/Biodiversity"},{"id":64471618,"name":"Earth and environmental sciences/Ocean sciences/Marine biology"},{"id":64471619,"name":"Scientific community and society/Scientific community/Policy"}],"tags":[],"updatedAt":"2026-04-09T08:13:08+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-09 08:13:08","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8894972","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8894972","identity":"rs-8894972","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}