A conceptual framework to help choose appropriate blue nature-based solutions.

Biodiversity loss and climate change have severely impacted ecosystems and livelihoods worldwide, compromising access to food and water, increasing disaster risk, and affecting human health globally. Nature-based Solutions (NbS) have gained interest in addressing these global societal challenges. Although much effort has been directed to NbS in urban and terrestrial environments, the implementation of NbS in marine and coastal environments (blue NbS) lags. The lack of a framework to guide decision-makers and practitioners through the initial planning stages appears to be one of the main obstacles to the slow implementation of blue NbS. To address this, we propose an integrated conceptual framework, built from expert knowledge, to inform the selection of the most appropriate blue NbS based on desired intervention objectives and social-ecological context. Our conceptual framework follows a four incremental steps structure: Step 1 aims to identify the societal challenge(s) to address; Step 2 highlights ecosystem services and the underlying biodiversity and ecological functions that could contribute to confronting the societal challenge(s); Step 3 identify the specific environmental context the intervention needs to be set within (e.g. the spatial scale the intervention will operate within, the ecosystem's vulnerability to stressors, and its ecological condition); and Step 4 provides a selection of potential blue NbS interventions that would help address the targeted societal challenge(s) considering the context defined through Step 3. Designed to maintain, enhance, recover, rehabilitate, or create ecosystem services by supporting biodiversity, the blue NbS intervention portfolio includes marine protection (i.e., fully, highly, lightly, and minimally protected areas), restorative activities (i.e., active, passive, and partial restoration; rehabilitation of ecological function and ecosystem creation), and other management measures (i.e., implementation and enforcement of regulation). Ultimately, our conceptual framework guides decision-makers toward a versatile portfolio of interventions that cater to the specific needs of each ecosystem rather than imposing a rigid, one-size-fits-all model. In the future, this framework needs to integrate socio-economic considerations more comprehensively and be kept up-to-date by including the latest scientific information.

The impact of mobile demersal fishing on carbon storage in seabed sediments.

Subtidal marine sediments are one of the planet's primary carbon stores and strongly influence the oceanic sink for atmospheric CO2 . By far the most widespread human activity occurring on the seabed is bottom trawling/dredging for fish and shellfish. A global first-order estimate suggested mobile demersal fishing activities may cause 0.16-0.4 Gt of organic carbon (OC) to be remineralized annually from seabed sediment carbon stores (Sala et al., 2021). There are, however, many uncertainties in this calculation. Here, we discuss the potential drivers of change in seabed sediment OC stores due to mobile demersal fishing activities and conduct a literature review, synthesizing studies where this interaction has been directly investigated. Under certain environmental settings, we hypothesize that mobile demersal fishing would reduce OC in seabed stores due to lower production of flora and fauna, the loss of fine flocculent material, increased sediment resuspension, mixing and transport and increased oxygen exposure. Reductions would be offset to varying extents by reduced faunal bioturbation and community respiration, increased off-shelf transport and increases in primary production from the resuspension of nutrients. Studies which directly investigated the impact of demersal fishing on OC stocks had mixed results. A finding of no significant effect was reported in 61% of 49 investigations; 29% reported lower OC due to fishing activities, with 10% reporting higher OC. In relation to remineralization rates within the seabed, four investigations reported that demersal fishing activities decreased remineralization, with three reporting higher remineralization rates. Patterns in the environmental and experimental characteristics between different outcomes were largely indistinct. More evidence is urgently needed to accurately quantify the impact of anthropogenic physical disturbance on seabed carbon in different environmental settings and to incorporate full evidence-based carbon considerations into global seabed management.

Marine reserves can mitigate and promote adaptation to climate change.

Strong decreases in greenhouse gas emissions are required to meet the reduction trajectory resolved within the 2015 Paris Agreement. However, even these decreases will not avert serious stress and damage to life on Earth, and additional steps are needed to boost the resilience of ecosystems, safeguard their wildlife, and protect their capacity to supply vital goods and services. We discuss how well-managed marine reserves may help marine ecosystems and people adapt to five prominent impacts of climate change: acidification, sea-level rise, intensification of storms, shifts in species distribution, and decreased productivity and oxygen availability, as well as their cumulative effects. We explore the role of managed ecosystems in mitigating climate change by promoting carbon sequestration and storage and by buffering against uncertainty in management, environmental fluctuations, directional change, and extreme events. We highlight both strengths and limitations and conclude that marine reserves are a viable low-tech, cost-effective adaptation strategy that would yield multiple cobenefits from local to global scales, improving the outlook for the environment and people into the future.

Addressing Criticisms of Large-Scale Marine Protected Areas.

Designated large-scale marine protected areas (LSMPAs, 100,000 or more square kilometers) constitute over two-thirds of the approximately 6.6% of the ocean and approximately 14.5% of the exclusive economic zones within marine protected areas. Although LSMPAs have received support among scientists and conservation bodies for wilderness protection, regional ecological connectivity, and improving resilience to climate change, there are also concerns. We identified 10 common criticisms of LSMPAs along three themes: (1) placement, governance, and management; (2) political expediency; and (3) social-ecological value and cost. Through critical evaluation of scientific evidence, we discuss the value, achievements, challenges, and potential of LSMPAs in these arenas. We conclude that although some criticisms are valid and need addressing, none pertain exclusively to LSMPAs, and many involve challenges ubiquitous in management. We argue that LSMPAs are an important component of a diversified management portfolio that tempers potential losses, hedges against uncertainty, and enhances the probability of achieving sustainably managed oceans.

Courtship and mating behaviour of manta rays Mobula alfredi and M. birostris in the Maldives.

The aim of this 14 year study was to elucidate the entire courtship and mating behaviour of manta rays Mobula alfredi and M. birostris using behavioural observations, video and photographic records. From 2003 to 2016, over 11,000 surveys were undertaken at known manta ray aggregation sites in the Maldives to record any observed manta rays reproductive activity. From 47,591 photo-ID sightings, 4,247 individual M. alfredi were identified and 226 individual M. birostris from 229 photo-ID sightings, all recorded at 22 atolls across 265 different sites. Courtship activity was observed on 206 surveys at 30 different sites. A total of 229 courtship events were recorded, with 90% (n = 205) of them occurring at cleaning sites. The observed courtship activity was categorised into seven distinct stages which are described in detail: initiation, endurance, evasion, pre-copulatory positioning, copulation, post-copulatory holding and separation. Photographs provide the first scientific record of the entirety of manta rays courtship and mating. Both M. alfredi and M. birostris appear to engage in the same elaborate courtship rituals, exhibiting the same behaviours during all stages of the courtship and mating process.

Identifying species at extinction risk using global models of anthropogenic impact.

The International Union for Conservation of Nature Red List of Endangered Species employs a robust, standardized approach to assess extinction threat focussed on taxa approaching an end-point in population decline. Used alone, we argue this enforces a reactive approach to conservation. Species not assessed as threatened but which occur predominantly in areas with high levels of anthropogenic impact may require proactive conservation management to prevent loss. We matched distribution and bathymetric range data from the global Red List assessment of 632 species of marine cone snails with human impacts and projected ocean thermal stress and aragonite saturation (a proxy for ocean acidification). Our results show 67 species categorized as 'Least Concern' have 70% or more of their occupancy in places subject to high and very high levels of human impact with 18 highly restricted species (range <100 km(2)) living exclusively in such places. Using a range-rarity scoring method we identified where clusters of endemic species are subject to all three stressors: high human impact, declining aragonite saturation levels and elevated thermal stress. Our approach reinforces Red List threatened status, highlights candidate species for reassessment, contributes important evidential data to minimize data deficiency and identifies regions and species for proactive conservation.

Climate change mitigation and nature conservation both require higher protected area targets.

Nations of the world have, to date, pursued nature protection and climate change mitigation and adaptation policies separately. Both efforts have failed to achieve the scale of action needed to halt biodiversity loss or mitigate climate change. We argue that success can be achieved by aligning targets for biodiversity protection with the habitat protection and restoration necessary to bring down greenhouse gas concentrations and promote natural and societal adaptation to climate change. Success, however, will need much higher targets for environmental protection than the present 10% of sea and 17% of land. A new target of 30% of the sea given high levels of protection from exploitation and harm by 2030 is under consideration and similar targets are being discussed for terrestrial habitats. We make the case here that these higher targets, if achieved, would make the transition to a warmer world slower and less damaging for nature and people. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

The timing and causality of ecological shifts on Caribbean reefs.

Caribbean reefs have experienced unprecedented changes in the past four decades. Of great concern is the perceived widespread shift from coral to macroalgal dominance and the question of whether it represents a new, stable equilibrium for coral-reef communities. The primary causes of the shift-grazing pressure (top-down), nutrient loading (bottom-up) or direct coral mortality (side-in)-still remain somewhat controversial in the coral-reef literature. We have attempted to tease out the relative importance of each of these causes. Four insights emerge from our analysis of an early regional dataset of information on the benthic composition of Caribbean reefs spanning the years 1977-2001. First, although three-quarters of reef sites have experienced coral declines concomitant with macroalgal increases, fewer than 10% of the more than 200 sites studied were dominated by macroalgae in 2001, by even the most conservative definition of dominance. Using relative dominance as the threshold, a total of 49 coral-to-macroalgae shifts were detected. This total represents ~35% of all sites that were dominated by coral at the start of their monitoring periods. Four shifts (8.2%) occurred because of coral loss with no change in macroalgal cover, 15 (30.6%) occurred because of macroalgal gain without coral loss, and 30 (61.2%) occurred owing to concomitant coral decline and macroalgal increase. Second, the timing of shifts at the regional scale is most consistent with the side-in model of reef degradation, which invokes coral mortality as a precursor to macroalgal takeover, because more shifts occurred after regional coral-mortality events than expected by chance. Third, instantaneous observations taken at the start and end of the time-series for individual sites showed these reefs existed along a continuum of coral and macroalgal cover. The continuous, broadly negative relationship between coral and macroalgal cover suggests that in some cases coral-to-macroalgae phase shifts may be reversed by removing sources of perturbation or restoring critical components such as the herbivorous sea urchin Diadema antillarum to the system. The five instances in which macroalgal dominance was reversed corroborate the conclusion that macroalgal dominance is not a stable, alternative community state as has been commonly assumed. Fourth, the fact that the loss in regional coral cover and concomitant changes to the benthic community are related to punctuated, discrete events with known causes (i.e. coral disease and bleaching), lends credence to the hypothesis that coral reefs of the Caribbean have been under assault from climate-change-related maladies since the 1970s.

Financial comparisons of fishing gear used in Kenya's coral reef lagoons.

The cost of fishing and the income earned by fishers using small and large traps, gill nets, beach seines, hand lines, and spearguns were assessed in the multigear fishery of southern Kenya to establish a financial rationale for fishing gear use. Direct observations and key-informant interviews with fish leaders and boat captains were used to gather data on fish catch, cost of fishing gear, boats, and the price of fish. Among the fishing gear used, spearguns had the lowest monthly cost (USD 1 mo(-1)) while big traps had the highest (USD 13 mo(-1)). Income was highest among capital cost beach seine fishers (USD 183 mo(-1)) and lowest among noncapital cost beach seine fishers (USD 20 mo(-1)). There was a direct positive correlation between income earned and profitability of gear. Correlation of the financial measure for each gear to four categories of damage to fish and habitats showed that low cost fishing gear were associated with the highest environmental damage indicating a trade-off between cost of gear and environmental health.

Conservation targets in marine protected area management suffer from shifting baseline syndrome: A case study on the Dogger Bank.

The Dogger Bank is a subtidal hill in the North Sea that is a candidate Special Area of Conservation under the EU Habitats Directive in UK waters. Historical records indicate that the Bank has been subject to human exploitation from before the 16th century but conservation objectives have been developed using recent survey data. This has the potential to significantly underestimate the alteration this ecosystem has experienced, making the Dogger Bank an example of shifting baseline syndrome in protected area management. We compile quantitative and qualitative descriptions from historical records of change in catch rates, fishing effort, price and fish size to show that there have been prolonged declines in abundance of fish on the Bank since the early 19th century. Use of present day data to inform conservation has led to unambitious recovery targets. Historical data, we argue, are an essential input to conservation decision making.

Diving behaviour of whale sharks in relation to a predictable food pulse.

We present diving data for four whale sharks in relation to a predictable food pulse (reef fish spawn) and an analysis of the longest continuous fine-resolution diving record for a planktivorous shark. Fine-resolution pressure data from a recovered pop-up archival satellite tag deployed for 206 days on a whale shark were analysed using the fast Fourier Transform method for frequency domain analysis of time-series. The results demonstrated that a free-ranging whale shark displays ultradian, diel and circa-lunar rhythmicity of diving behaviour. Whale sharks dive to over 979.5 m and can tolerate a temperature range of 26.4 degrees C. The whale sharks made primarily diurnal deep dives and remained in relatively shallow waters at night. Whale shark diving patterns are influenced by a seasonally predictable food source, with shallower dives made during fish spawning periods.

Public awareness and attitudes towards marine protection in the United Kingdom.

Public perception research evaluating awareness and attitudes towards marine protection is limited in the United Kingdom (UK) and worldwide. Given public opinion can help drive policy and affect its successful delivery we conducted nationwide surveys in 2005, 2010 and 2015 to assess public knowledge of UK (England, Scotland and Wales) sea 'health' and management. Respondents from all three surveys were relatively pessimistic about sea 'health', perceiving this as poor-fair and largely in decline. Enthusiasm for marine conservation was high with almost two-thirds of respondents in each survey wanting >40% of UK seas highly protected from fishing and damaging activities. In 2015 there was considerable dissatisfaction with the rate of progress in Marine Conservation Zone designation and over three-quarters of respondents considered dredging and trawling to be inappropriate in protected areas, contrary to management. The UK government and devolved administrations need to better align future conservation and management with public expectations.

Rapidly shifting environmental baselines among fishers of the Gulf of California.

Shifting environmental baselines are inter-generational changes in perception of the state of the environment. As one generation replaces another, people's perceptions of what is natural change even to the extent that they no longer believe historical anecdotes of past abundance or size of species. Although widely accepted, this phenomenon has yet to be quantitatively tested. Here we survey three generations of fishers from Mexico's Gulf of California (N=108), where fish populations have declined steeply over the last 60 years, to investigate how far and fast their environmental baselines are shifting. Compared to young fishers, old fishers named five times as many species and four times as many fishing sites as once being abundant/productive but now depleted (Kruskal-Wallis tests, both p<0.001) with no evidence of a slowdown in rates of loss experienced by younger compared to older generations (Kruskal-Wallis test, n.s. in both cases). Old fishers caught up to 25 times as many Gulf grouper Mycteroperca jordani as young fishers on their best ever fishing day (regression r(2)=0.62, p<0.001). Despite times of plentiful large fish still being within living memory, few young fishers appreciated that large species had ever been common or nearshore sites productive. Such rapid shifts in perception of what is natural help explain why society is tolerant of the creeping loss of biodiversity. They imply a large educational hurdle in efforts to reset expectations and targets for conservation.

Sharks and people: insight into the global practices of tourism operators and their attitudes to shark behaviour.

Shark tourism is a popular but controversial activity. We obtained insights into this industry via a global e-mailed questionnaire completed by 45 diving/snorkelling operators who advertised shark experiences (shark operators) and 49 who did not (non-shark operators). 42% of shark operators used an attractant to lure sharks and 93% stated they had a formal code of conduct which 86% enforced "very strictly". While sharks were reported to normally ignore people, 9 operators had experienced troublesome behaviour from them. Whilst our research corroborates previous studies indicating minimal risk to humans from most shark encounters, a precautionary approach to provisioning is required to avoid potential ecological and societal effects of shark tourism. Codes of conduct should always stipulate acceptable diver behaviour and appropriate diver numbers and shark operators should have a moral responsibility to educate their customers about the need for shark conservation.

Sessile and mobile components of a benthic ecosystem display mixed trends within a temperate marine reserve.

Despite recent efforts to increase the global coverage of marine protected areas (MPAs), studies investigating the effectiveness of marine protected areas within temperate waters remain scarce. Furthermore, out of the few studies published on MPAs in temperate waters, the majority focus on specific ecological or fishery components rather than investigating the ecosystem as a whole. This study therefore investigated the dynamics of both benthic communities and fish populations within a recently established, fully protected marine reserve in Lamlash Bay, Isle of Arran, United Kingdom, over a four year period. A combination of photo and diver surveys revealed live maerl (Phymatolithon calcareum), macroalgae, sponges, hydroids, feather stars and eyelash worms (Myxicola infundibulum) to be significantly more abundant within the marine reserve than on surrounding fishing grounds. Likewise, the overall composition of epifaunal communities in and outside the reserve was significantly different. Both results are consistent with the hypothesis that protecting areas from fishing can encourage seafloor habitats to recover. In addition, the greater abundance of complex habitats within the reserve appeared to providing nursery habitat for juvenile cod (Gadus morhua) and scallops (Pecten maximus and Aequipecten opercularis). In contrast, there was little difference in the abundance of mobile benthic fauna, such as crabs and starfish, between the reserve and outside. Similarly, the use of baited underwater video cameras revealed no difference in the abundance and size of fish between the reserve and outside. Limited recovery of these ecosystem components may be due to the relatively small size (2.67 km(2)) and young age of the reserve (<5 years), both of which might have limited the extent of any benefits afforded to mobile fauna and fish communities. Overall, this study provides evidence that fully protected marine reserves can encourage seafloor habitats to recover, which in turn, can create a number of benefits that flow back to other species, including those of commercial importance.

The origins of intensive marine fishing in medieval Europe: the English evidence.

The catastrophic impact of fishing pressure on species such as cod and herring is well documented. However, the antiquity of their intensive exploitation has not been established. Systematic catch statistics are only available for ca.100 years, but large-scale fishing industries existed in medieval Europe and the expansion of cod fishing from the fourteenth century (first in Iceland, then in Newfoundland) played an important role in the European colonization of the Northwest Atlantic. History has demonstrated the scale of these late medieval and post-medieval fisheries, but only archaeology can illuminate earlier practices. Zooarchaeological evidence shows that the clearest changes in marine fishing in England between AD 600 and 1600 occurred rapidly around AD 1000 and involved large increases in catches of herring and cod. Surprisingly, this revolution predated the documented post-medieval expansion of England's sea fisheries and coincided with the Medieval Warm Period--when natural herring and cod productivity was probably low in the North Sea. This counterintuitive discovery can be explained by the concurrent rise of urbanism and human impacts on freshwater ecosystems. The search for 'pristine' baselines regarding marine ecosystems will thus need to employ medieval palaeoecological proxies in addition to recent fisheries data and early modern historical records.

Effects of Fishing on the Ecosystem Structure of Coral Reefs.

Overfishing is considered one of the three most significant threats to coral reef ecosystems. Exponentially increasing human populations in the tropics have placed enormous demands upon reefs as a food source. At high intensities, termed ecosystem or Malthusian overfishing, fishing causes major direct and indirect effects on the community structure of fishes and other organisms. It reduces species diversity and leads to local extinctions not only of target species but also of other species not fished directly. Conceivably it could also lead to global extinctions. Loss of keystone species, such as predators of echinoderms, through fishing, can lead to major effects on reef processes, such as accretion of calcium carbonate. Ultimately, sustained heavy fishing may lead to loss of entire functional groups of species, resulting in impairment of the potentially important ecosystem functions provided by those groups. Overfishing has been shown to interact with other agents of disturbance to reduce the ability of reefs to recover from natural occurrences such as hurricanes. Effective management of fishing will require a deeper understanding of the effects of exploitation than we now possess. Research initiatives are underway to examine the responses of fish populations to fishing, generally responses to protection from fishing. There is, however, an urgent need to look beyond fish communities and to consider the entire reef ecosystem. Studies that integrate population and community biology with ecosystem processes will provide a much better understanding of the effects of biodiversity loss on reef function and will improve our ability to manage these complex systems. Efecto de la pesca sobre la estructura ecosistémica de los arrecifes de coral.

Resumen: La sobrepesca es considerada como una de las tres amenazas más significativas para los arrecifes de coral. En los trópicos, las poblaciones humanas en incremento exponencial han impuesto demandas enormes sobre los arrecifes como fuentes de alimentos. A intensidades de pesca altas, denominadas sobrepesca ecosistémica o Maltusiana, la pesca causa importantes efectos directos e indirectos en la estructura de la comunidad de peces y otros organismos. La sobrepesca reduce la diversidad de especies y lleva a la extinción local tanto de las especies pescadas como de otras especies no pescadas en forma directa. En forma concebible puede también llevar a extinciones globales. La pérdida de especies clave por medio de la pesca, tales como predadores de equinodermos, puede tener ejectos importantes sobre los procesos biológicos del arrecife, tales como la acumulación de carbonato de calcio. En última instancia, la pesca intensiva sostenida puede llever a la pérdida de grupos funcionales de especies enteras, lo que resulta en la pédida de las funciones ecosistémicas potencialmente importantes provistas por estas especies. Se ha demostrado que la sobrepesca interactúa con otros agentes perturbadores para reducir la habilidad de los arrecifes para recuperarse de fenómenos naturales como los huracanes. El manejo pesquero efectivo requerirá de un entendimiento más profundo de los efectos de la explotación. Se están llevando a cabo iniciativas de investigación para examinar las respuestas de las poblaciones de peces a la pesca, se trata generalmente de estudios sobre la respuesta de las poblaciones a la protección de la pesca. Existe sin embargo, una necesidad urgente de mirar más alla de las comunidades de peces y considerar a la totalidad del ecosistema. Los estudios que integran la biología de poblaciones y comunidades con los procesos ecosistémicos, proveerián de un mejor entendimiento de los efectos de la pérdida de la biodiversidad sobre las funciones de los arrecifes y mejorarián nuestra habilidad para manejar estos sistemas complejos.

Partial mortality in massive reef corals as an indicator of sediment stress on coral reefs.

Partial mortality and fission on colonies of four common massive coral species were examined at sites differing in their exposure to river sediments in St. Lucia, West Indies. Rates of partial mortality were higher close to the river mouths, where more sediments were deposited, than away from the rivers in two coral species. Frequency of fission showed no significant trend. The percent change in coral cover on reefs from 1995 to 1998 was negatively related to the rate of partial mortality estimated in 1998 in all species. This suggests that partial mortality rates could reflect longer-term temporal changes in coral communities. Similar conclusions could also be reached using a less precise measure and simply recording partial mortality on colonies as <50% and >/=50% dead tissue. We conclude that partial mortality in some species of massive reef corals, expressed as the amount of dead tissue per colony, could provide a rapid and effective means of detecting sediment stress on coral reefs.

The past and future of fish consumption: Can supplies meet healthy eating recommendations?

In many developed countries fish and shellfish are increasingly promoted as healthy alternatives to other animal protein. We analysed how much fish was available to UK and global populations after accounting for processing losses, and compared this to recommended levels of fish consumption. In 2012, UK domestic fish landings per capita fell 81% below the recommended intake, although declines were masked by increased imports and aquaculture from the 1970s onwards. Global wild fish supply per capita declined by 32% from its peak in 1970. However, overall fish supplies per capita increased by 10% over the same period due to rapidly expanding aquaculture production. Whilst aquaculture has so far prevented a downturn in global fish supplies, many developed nations continue to aspire to consume more fish than they produce. Until demand is balanced with sustainable methods of production governments should consider carefully the social and environmental implications of greater fish consumption.

The contribution of very large marine protected areas to marine conservation: giant leaps or smoke and mirrors?

In recent years, marine protected areas have been "super-sized". At first glance, this seems a gift to marine conservation. Yet, the new wave of very large marine protected areas ("VLMPAs") have faced criticism from the scientific community. In this article we examine the merits and the criticisms of VLMPAS, and consider whether they provide a much-needed boost to marine conservation, or are simply too good to be true.