A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014–2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.     

Low genetic diversity in a marine nature reserve: re-evaluating diversity criteria in reserve design

Little consideration has been given to the genetic composition of populations associated with marine reserves, as reserve designation is generally to protect specific species, communities or habitats. Nevertheless, it is important to conserve genetic diversity since it provides the raw material for the maintenance of species diversity over longer, evolutionary time-scales and may also confer the basis for adaptation to environmental change. Many current marine reserves are small in size and isolated to some degree (e.g. sea loughs and offshore islands). While such features enable easier management, they may have important implications for the genetic structure of protected populations, the ability of populations to recover from local catastrophes and the potential for marine reserves to act as sources of propagules for surrounding areas. Here, we present a case study demonstrating genetic differentiation, isolation, inbreeding and reduced genetic diversity in populations of the dogwhelk Nucella lapillus in Lough Hyne Marine Nature Reserve (an isolated sea lough in southern Ireland), compared with populations on the local adjacent open coast and populations in England, Wales and France. Our study demonstrates that this sea lough is isolated from open coast populations, and highlights that there may be long-term genetic consequences of selecting reserves on the basis of isolation and ease of protection.     

Ecosystem‐based management of coral reefs under climate change

Coral reefs provide food and livelihoods for hundreds of millions of people as well as harbour some of the highest regions of biodiversity in the ocean. However, overexploitation, land‐use change and other local anthropogenic threats to coral reefs have left many degraded. Additionally, coral reefs are faced with the dual emerging threats of ocean warming and acidification due to rising CO₂ emissions, with dire predictions that they will not survive the century. This review evaluates the impacts of climate change on coral reef organisms, communities and ecosystems, focusing on the interactions between climate change factors and local anthropogenic stressors. It then explores the shortcomings of existing management and the move towards ecosystem‐based management and resilience thinking, before highlighting the need for climate change‐ready marine protected areas (MPAs), reduction in local anthropogenic stressors, novel approaches such as human‐assisted evolution and the importance of sustainable socialecological systems. It concludes that designation of climate change‐ready MPAs, integrated with other management strategies involving stakeholders and participation at multiple scales such as marine spatial planning, will be required to maximise coral reef resilience under climate change. However, efforts to reduce carbon emissions are critical if the long‐term efficacy of local management actions is to be maintained and coral reefs are to survive.     

Climate warming, marine protected areas and the ocean-scale integrity of coral reef ecosystems

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.     

Marine protected areas can be useful but are not a silver bullet for kelp conservation

Kelp forests are among the most valuable ecosystems on Earth, but they are increasingly being degraded and lost due to a range of human-related stressors, leading to recent calls for their improved management and conservation. One of the primary tools to conserve marine species and biodiversity is the establishment of marine protected areas (MPAs). International commitments to protect 30% of the world's ecosystems are gaining momentum, offering a promising avenue to secure kelp forests into the Anthropocene. However, a clear understanding of the efficacy of MPAs for conserving kelp forests in a changing ocean is lacking. In this perspective, we question whether strengthened global protection will create meaningful conservation outcomes for kelp forests. We explore the benefits of MPAs for kelp conservation under a suite of different stressors, focusing on empirical evidence from protected kelp forests. We show that MPAs can be effective against some drivers of kelp loss (e.g., overgrazing, kelp harvesting), particularly when they are maintained in the long-term and enforced as no-take areas. There is also some evidence that MPAs can reduce impacts of climate change through building resilience in multi-stressor situations. However, MPAs also often fail to provide protection against ocean warming, marine heatwaves, coastal darkening, and pollution, which have emerged as dominant drivers of kelp forest loss globally. Although well-enforced MPAs should remain an important tool to protect kelp forests, successful kelp conservation will require implementing an additional suite of management solutions that target these accelerating threats.     

The role of marine reserves in achieving sustainable fisheries

Many fishery management tools currently in use have conservation value. They are designed to maintain stocks of commercially important species above target levels. However, their limitations are evident from continuing declines in fish stocks throughout the world. We make the case that to reverse fishery declines, safeguard marine life and sustain ecosystem processes, extensive marine reserves that are off limits to fishing must become part of the management strategy. Marine reserves should be incorporated into modern fishery management because they can achieve many things that conventional tools cannot. Only complete and permanent protection from fishing can protect the most sensitive habitats and vulnerable species. Only reserves will allow the development of natural, extended age structures of target species, maintain their genetic variability and prevent deleterious evolutionary change from the effects of fishing. Species with natural age structures will sustain higher rates of reproduction and will be more resilient to environmental variability. Higher stock levels maintained by reserves will provide insurance against management failure, including risk-prone quota setting, provided the broader conservation role of reserves is firmly established and legislatively protected. Fishery management measures outside protected areas are necessary to complement the protection offered by marine reserves, but cannot substitute for it.     

Moving forward in global‐change ecology: capitalizing on natural variability

Natural resources managers are being asked to follow practices that accommodate for the impact of climate change on the ecosystems they manage, while global‐ecosystems modelers aim to forecast future responses under different climate scenarios. However, the lack of scientific knowledge about short‐term ecosystem responses to climate change has made it difficult to define set conservation practices or to realistically inform ecosystem models. Until recently, the main goal for ecologists was to study the composition and structure of communities and their implications for ecosystem function, but due to the probable magnitude and irreversibility of climate‐change effects (species extinctions and loss of ecosystem function), a shorter term focus on responses of ecosystems to climate change is needed. We highlight several underutilized approaches for studying the ecological consequences of climate change that capitalize on the natural variability of the climate system at different temporal and spatial scales. For example, studying organismal responses to extreme climatic events can inform about the resilience of populations to global warming and contribute to the assessment of local extinctions. Translocation experiments and gene expression are particular useful to quantitate a species' acclimation potential to global warming. And studies along environmental gradients can guide habitat restoration and protection programs by identifying vulnerable species and sites. These approaches identify the processes and mechanisms underlying species acclimation to changing conditions, combine different analytical approaches, and can be used to improve forecasts of the short‐term impacts of climate change and thus inform conservation practices and ecosystem models in a meaningful way.     

The potential impacts of global climate change on marine protected areas

The potential effects of global climate changeon marine protected areas do not appear to havebeen addressed in the literature. This paperexamines the literature on protected areas,conservation biology, marine ecology,oceanography, and climate change, and reviewssome of the relevant differences between marineand terrestrial environments. Frameworks andclassifications systems used in protected areadesign are discussed. Finally, a frameworkthat summarizes some of the importantoceanographic processes and their links to thefood chain are reviewed. Species abundance anddistribution are expected to change as a resultof global climate change, potentiallycompromising the efficacy of marine protectedareas as biodiversity conservation tools. Thisreview suggests the need for: furtherinterdisciplinary research and the use oflinked models; an increase in marine protectedareas for biodiversity conservation and asresearch sites for teasing apart fishingeffects from climate effects; a temporallyresponsive approach to siting new marineprotected areas, shifting their locations ifnecessary; and large-scale ecosystem/integratedmanagement approaches to address the competinguses of the oceans and boundary-less threatssuch as global climate change and pollution.     

Climate Change or Land Use Dynamics: Do We Know What Climate Change Indicators Indicate?

Different components of global change can have interacting effects on biodiversity and this may influence our ability to detect the specific consequences of climate change through biodiversity indicators. Here, we analyze whether climate change indicators can be affected by land use dynamics that are not directly determined by climate change. To this aim, we analyzed three community-level indicators of climate change impacts that are based on the optimal thermal environment and average latitude of the distribution of bird species present at local communities. We used multiple regression models to relate the variation in climate change indicators to: i) environmental temperature; and ii) three landscape gradients reflecting important current land use change processes (land abandonment, fire impacts and urbanization), all of them having forest areas at their positive extremes. We found that, with few exceptions, landscape gradients determined the figures of climate change indicators as strongly as temperature. Bird communities in forest habitats had colder-dwelling bird species with more northern distributions than farmland, burnt or urban areas. Our results show that land use changes can reverse, hide or exacerbate our perception of climate change impacts when measured through community-level climate change indicators. We stress the need of an explicit incorporation of the interactions between climate change and land use dynamics to understand what are current climate change indicators indicating and be able to isolate real climate change impacts.     

Decadal-scale rebuilding of predator biomass in Philippine marine reserves

No-take marine reserves (NTMRs) provide hope that local carrying capacity may be partially restored if reserves are protected long enough. How long is long enough? We assess the duration of protection required for populations of large predatory reef fish in marine reserves to attain new steady states. We monitored biomass of large predatory fish in two marine reserves at Sumilon and Apo Islands, Philippines, almost annually for 26 years (1983–2009), and fit a logistic model to the data. As duration of reserve protection increased, biomass of predatory fish approached an asymptote, although the models suggest that 20–40 years of protection is required to attain new steady states. Thus, for local carrying capacity to be rebuilt, no-take protection must be effective on decadal timescales.     

Animal behaviour shapes the ecological effects of ocean acidification and warming: moving from individual to community‐level responses

Biological communities are shaped by complex interactions between organisms and their environment as well as interactions with other species. Humans are rapidly changing the marine environment through increasing greenhouse gas emissions, resulting in ocean warming and acidification. The first response by animals to environmental change is predominantly through modification of their behaviour, which in turn affects species interactions and ecological processes. Yet, many climate change studies ignore animal behaviour. Furthermore, our current knowledge of how global change alters animal behaviour is mostly restricted to single species, life phases and stressors, leading to an incomplete view of how coinciding climate stressors can affect the ecological interactions that structure biological communities. Here, we first review studies on the effects of warming and acidification on the behaviour of marine animals. We demonstrate how pervasive the effects of global change are on a wide range of critical behaviours that determine the persistence of species and their success in ecological communities. We then evaluate several approaches to studying the ecological effects of warming and acidification, and identify knowledge gaps that need to be filled, to better understand how global change will affect marine populations and communities through altered animal behaviours. Our review provides a synthesis of the far‐reaching consequences that behavioural changes could have for marine ecosystems in a rapidly changing environment. Without considering the pervasive effects of climate change on animal behaviour we will limit our ability to forecast the impacts of ocean change and provide insights that can aid management strategies.     

Evidence of artisanal fishing impacts and depth refuge in assemblages of Fijian reef fish

Protection from fishing generally results in an increase in the abundance and biomass of species targeted by fisheries within marine reserve boundaries. Natural refuges such as depth may also protect such species, yet few studies in the Indo Pacific have investigated the effects of depth concomitant with marine reserves. We studied the effects of artisanal fishing and depth on reef fish assemblages in the Kubulau District of Vanua Levu Island, Fiji, using baited remote underwater stereo-video systems. Video samples were collected from shallow (5–8 m) and deep (25–30 m) sites inside and outside of a large old marine reserve (60.6 km², 13 years old) and a small new marine reserve (4.25 km², 4 years old). Species richness tended to be greater in the shallow waters of the large old reserve when compared to fished areas. In the deeper waters, species richness appeared to be comparable. The difference in shallow waters was driven by species targeted by fisheries, indicative of a depth refuge effect. In contrast, differences in the abundance composition of the fish assemblage existed between protected and fished areas for deep sites, but not shallow. Fish species targeted by local fisheries were 89% more abundant inside the large old reserve than surrounding fished areas, while non-targeted species were comparable. We observed no difference in the species richness or abundance of species targeted by fisheries inside and outside of the small new reserve. This study suggests that artisanal fishing impacts on the abundance and species richness of coral reef fish assemblages and effects of protection are more apparent with large reserves that have been established for a long period of time. Observed effects of protection also vary with depth, highlighting the importance of explicitly incorporating multiple depth strata in studies of marine reserves.     

Vulnerability of the global terrestrial ecosystems to climate change

Climate change has far‐reaching impacts on ecosystems. Recent attempts to quantify such impacts focus on measuring exposure to climate change but largely ignore ecosystem resistance and resilience, which may also affect the vulnerability outcomes. In this study, the relative vulnerability of global terrestrial ecosystems to short‐term climate variability was assessed by simultaneously integrating exposure, sensitivity, and resilience at a high spatial resolution (0.05°). The results show that vulnerable areas are currently distributed primarily in plains. Responses to climate change vary among ecosystems and deserts and xeric shrublands are the most vulnerable biomes. Global vulnerability patterns are determined largely by exposure, while ecosystem sensitivity and resilience may exacerbate or alleviate external climate pressures at local scales; there is a highly significant negative correlation between exposure and sensitivity. Globally, 61.31% of the terrestrial vegetated area is capable of mitigating climate change impacts and those areas are concentrated in polar regions, boreal forests, tropical rainforests, and intact forests. Under current sensitivity and resilience conditions, vulnerable areas are projected to develop in high Northern Hemisphere latitudes in the future. The results suggest that integrating all three aspects of vulnerability (exposure, sensitivity, and resilience) may offer more comprehensive and spatially explicit adaptation strategies to reduce the impacts of climate change on terrestrial ecosystems.     

Interdisciplinary knowledge exchange across scales in a globally changing marine environment

The effects of anthropogenic global environmental change on biotic and abiotic processes have been reported in aquatic systems across the world. Complex synergies between concurrent environmental stressors and the resilience of the system to regime shifts, which vary in space and time, determine the capacity for marine systems to maintain structure and function with global environmental change. Consequently, an interdisciplinary approach that facilitates the development of new methods for the exchange of knowledge between scientists across multiple scales is required to effectively understand, quantify and predict climate impacts on marine ecosystem services. We use a literature review to assess the limitations and assumptions of current pathways to exchange interdisciplinary knowledge and the transferability of research findings across spatial and temporal scales and levels of biological organization to advance scientific understanding of global environmental change in marine systems. We found that species‐specific regional scale climate change research is most commonly published, and “supporting” is the ecosystem service most commonly referred to in publications. In addition, our paper outlines a trajectory for the future development of integrated climate change science for sustaining marine ecosystem services such as investment in interdisciplinary education and connectivity between disciplines.     

Scale‐dependent complementarity of climatic velocity and environmental diversity for identifying priority areas for conservation under climate change

As most regions of the earth transition to altered climatic conditions, new methods are needed to identify refugia and other areas whose conservation would facilitate persistence of biodiversity under climate change. We compared several common approaches to conservation planning focused on climate resilience over a broad range of ecological settings across North America and evaluated how commonalities in the priority areas identified by different methods varied with regional context and spatial scale. Our results indicate that priority areas based on different environmental diversity metrics differed substantially from each other and from priorities based on spatiotemporal metrics such as climatic velocity. Refugia identified by diversity or velocity metrics were not strongly associated with the current protected area system, suggesting the need for additional conservation measures including protection of refugia. Despite the inherent uncertainties in predicting future climate, we found that variation among climatic velocities derived from different general circulation models and emissions pathways was less than the variation among the suite of environmental diversity metrics. To address uncertainty created by this variation, planners can combine priorities identified by alternative metrics at a single resolution and downweight areas of high variation between metrics. Alternately, coarse‐resolution velocity metrics can be combined with fine‐resolution diversity metrics in order to leverage the respective strengths of the two groups of metrics as tools for identification of potential macro‐ and microrefugia that in combination maximize both transient and long‐term resilience to climate change. Planners should compare and integrate approaches that span a range of model complexity and spatial scale to match the range of ecological and physical processes influencing persistence of biodiversity and identify a conservation network resilient to threats operating at multiple scales.     

Marine protected areas increase resilience among coral reef communities

With marine biodiversity declining globally at accelerating rates, maximising the effectiveness of conservation has become a key goal for local, national and international regulators. Marine protected areas (MPAs) have been widely advocated for conserving and managing marine biodiversity yet, despite extensive research, their benefits for conserving non‐target species and wider ecosystem functions remain unclear. Here, we demonstrate that MPAs can increase the resilience of coral reef communities to natural disturbances, including coral bleaching, coral diseases, Acanthaster planci outbreaks and storms. Using a 20‐year time series from Australia's Great Barrier Reef, we show that within MPAs, (1) reef community composition was 21–38% more stable; (2) the magnitude of disturbance impacts was 30% lower and (3) subsequent recovery was 20% faster that in adjacent unprotected habitats. Our results demonstrate that MPAs can increase the resilience of marine communities to natural disturbance possibly through herbivory, trophic cascades and portfolio effects.     

Potential loss of genetic variability despite well established network of reserves: the case of the Iberian endemic lizard <Emphasis Type="Italic">Lacerta schreiberi</Emphasis>

Although future anthropogenic climate change is recognized as one of the major threats to European species, its implementation during reserve planning has only been started recently. We here describe climate change impacts on the Iberian endemic lizard Lacerta schreiberi expecting serious declines and range reductions due to a loss of suitable climate space in the next future. We apply species distribution models to assess possible future changes in the lizard’s range, identify areas with high extinction risk meriting conservation efforts and analyze whether the Natura 2000 network in its current stage will offer a sufficient protection for the genetically most valuable lineages. Despite a very good coverage and connectivity of the most valuable populations of L. schreiberi with the existing protected sites network, our results predict a strong loss of genetic variability by 2080. Also, two main patterns become evident: While the genetically less diverse north-western populations may be less affected by climate change, the climate change effects on the southern isolates and the genetically most diverse populations within the Central System may be devastating. To improve a successful prospective conservation of L. schreiberi the management of protected sites needs to consider the processes that threaten this species. Furthermore, our study highlights the urgent need to consider climate change effects on evolutionary significant units within the Natura 2000 framework.     

It is getting hotter in here: determining and projecting the impacts of global environmental change on drylands

Drylands occupy large portions of the Earth, and are a key terrestrial biome from the socio-ecological point of view. In spite of their extent and importance, the impacts of global environmental change on them remain poorly understood. In this introduction, we review some of the main expected impacts of global change in drylands, quantify research efforts on the topic, and highlight how the articles included in this theme issue contribute to fill current gaps in our knowledge. Our literature analyses identify key under-studied areas that need more research (e.g. countries such as Mauritania, Mali, Burkina Faso, Chad and Somalia, and deserts such as the Thar, Kavir and Taklamakan), and indicate that most global change research carried out to date in drylands has been done on a unidisciplinary basis. The contributions included here use a wide array of organisms (from micro-organisms to humans), spatial scales (from local to global) and topics (from plant demography to poverty alleviation) to examine key issues to the socio-ecological impacts of global change in drylands. These papers highlight the complexities and difficulties associated with the prediction of such impacts. They also identify the increased use of long-term experiments and multidisciplinary approaches as priority areas for future dryland research. Major advances in our ability to predict and understand global change impacts on drylands can be achieved by explicitly considering how the responses of individuals, populations and communities will in turn affect ecosystem services. Future research should explore linkages between these responses and their effects on water and climate, as well as the provisioning of services for human development and well-being.     

Signals of Climate Change in Butterfly Communities in a Mediterranean Protected Area

The European protected-area network will cease to be efficient for biodiversity conservation, particularly in the Mediterranean region, if species are driven out of protected areas by climate warming. Yet, no empirical evidence of how climate change influences ecological communities in Mediterranean nature reserves really exists. Here, we examine long-term (1998–2011/2012) and short-term (2011–2012) changes in the butterfly fauna of Dadia National Park (Greece) by revisiting 21 and 18 transects in 2011 and 2012 respectively, that were initially surveyed in 1998. We evaluate the temperature trend for the study area for a 22-year-period (1990–2012) in which all three butterfly surveys are included. We also assess changes in community composition and species richness in butterfly communities using information on (a) species’ elevational distributions in Greece and (b) Community Temperature Index (calculated from the average temperature of species'' geographical ranges in Europe, weighted by species'' abundance per transect and year). Despite the protected status of Dadia NP and the subsequent stability of land use regimes, we found a marked change in butterfly community composition over a 13 year period, concomitant with an increase of annual average temperature of 0.95°C. Our analysis gave no evidence of significant year-to-year (2011–2012) variability in butterfly community composition, suggesting that the community composition change we recorded is likely the consequence of long-term environmental change, such as climate warming. We observe an increased abundance of low-elevation species whereas species mainly occurring at higher elevations in the region declined. The Community Temperature Index was found to increase in all habitats except agricultural areas. If equivalent changes occur in other protected areas and taxonomic groups across Mediterranean Europe, new conservation options and approaches for increasing species’ resilience may have to be devised.     

Ocean acidification refugia in variable environments

Climate change refugia in the terrestrial biosphere are areas where species are protected from global environmental change and arise from natural heterogeneity in landscapes and climate. Within the marine realm, ocean acidification, or the global decline in seawater pH, remains a pervasive threat to organisms and ecosystems. Natural variability in seawater carbon dioxide (CO₂) chemistry, however, presents an opportunity to identify ocean acidification refugia (OAR) for marine species. Here, we review the literature to examine the impacts of variable CO₂ chemistry on biological responses to ocean acidification and develop a framework of definitions and criteria that connects current OAR research to management goals. Under the concept of managing vulnerability, the most likely mechanisms by which OAR can mitigate ocean acidification impacts are by reducing exposure to harmful conditions or enhancing adaptive capacity. While local management options, such as OAR, show some promise, they present unique challenges, and reducing global anthropogenic CO₂ emissions must remain a priority.