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Healthy marine ecosystems provide critical benefits to people worldwide, but increasing threats from climate change and human activities disrupt ecosystem functionality and put these benefits at risk. Local and regional assessments have shown these impacts can be substantial, but we lack a global assessment of risk to marine biodiversity. Here we assessed risk of impact by intersecting spatial distributions of 21,267 marine animal species with distributions of 13 anthropogenic stressors according to each species’ vulnerability, examining results through multiple lenses that connect to different conservation objectives: species, taxon, and functional vulnerability. Using this species-focused approach, we found that vulnerable functional entities in coastal ecosystems across all marine ecological provinces are at greater risk of impact than indicated by assessments of broader ecosystem-level impact risk based on vulnerability of representative habitats, driven largely by climate stressors. Where multiple lenses of impact assessment indicate elevated risk, broad area-based protections may be warranted, but where impacts are focused on vulnerable functional entities there may be opportunities for more narrowly targeted conservation strategies such as local habitat restoration, assisted migration, or fishing gear restrictions. These results provide key insights at local to global scales on where and how to best meet conservation of species diversity and ecosystem function.

Marine species are declining at an unprecedented rate, catalyzing many nations to adopt conservation and management targets within their jurisdictions. However, marine species and the biophysical processes that sustain them are naive to international borders. An understanding of the prevalence of cross-border species distributions is important for informing high-level conservation strategies, such as bilateral or regional agreements. Here, we examined 28,252 distribution maps to determine the number and locations of transboundary marine plants and animals. More than 90% of species have ranges spanning at least two jurisdictions, with 58% covering more than 10 jurisdictions. All jurisdictions have at least one transboundary species, with the highest concentrations of transboundary species in the USA, Australia, Indonesia, and the Areas Beyond National Jurisdiction. Distributions of mapped biodiversity indicate that overcoming the challenges of multinational governance is critical for a much wider suite of species than migratory megavertebrates and commercially exploited fish stocks-the groups that have received the vast majority of multinational management attention. To effectively protect marine biodiversity, international governance mechanisms (particularly those related to the Convention on Biological Diversity, the Convention on Migratory Species, and Regional Seas Organizations) must be expanded to promote multinational conservation planning, and complimented by a holistic governance framework for biodiversity beyond national jurisdiction.

Feeding a growing, increasingly affluent population while limiting environmental pressures of food production is a central challenge for society. Understanding the location and magnitude of food production is key to addressing this challenge because pressures vary substantially across food production types. Applying data and models from life cycle assessment with the methodologies for mapping cumulative environmental impacts of human activities (hereafter cumulative impact mapping) provides a powerful approach to spatially map the cumulative environmental pressure of food production in a way that is consistent and comprehensive across food types. However, these methodologies have yet to be combined. By synthesizing life cycle assessment and cumulative impact mapping methodologies, we provide guidance for comprehensively and cumulatively mapping the environmental pressures (e.g., greenhouse gas emissions, spatial occupancy, and freshwater use) associated with food production systems. This spatial approach enables quantification of current and potential future environmental pressures, which is needed for decision makers to create more sustainable food policies and practices. ª 2020 The Author(s). Published by Elsevier Inc. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

Management of the land-sea interface is essential for global conservation and sustainability objectives because coastal regions maintain natural processes that support biodiversity and the livelihood of billions of people. However, assessments of coastal regions have focused strictly on either the terrestrial or marine realm. Consequently, understanding of the overall state of Earth's coastal regions is poor. We integrated the terrestrial human footprint and marine cumulative human impact maps in a global assessment of the anthropogenic pressures affecting coastal regions. Of coastal regions globally, 15.5% had low anthropogenic pressure, mostly in Canada, Russia, and Greenland. Conversely, 47.9% of coastal regions were heavily affected by humanity, and in most countries (84.1%)50% of their coastal regions were degraded. Nearly half (43.3%) of protected areas across coastal regions were exposed to high human pressures. To meet global sustainability objectives, all nations must undertake greater actions to preserve and restore the coastal regions within their borders.costa, huella humana, impacto humano cumulativo, litoral, presión humana, restauración, tierras vírgenes Resumen El manejo de la interfaz entre la tierra y el mar es esencial para los objetivos mundiales de conservación y sustentabilidad ya que las regiones costeras mantienen los procesos naturales que sostienen a la biodiversidad y al sustento de miles de millones de personas. Sin embargo, los análisis de las regiones costeras se han enfocado estrictamente en el ámbito marino o en el terrestre, pero no en ambos. Por consiguiente, el conocimiento del estado general de las regiones costeras del planeta es muy pobre. Integramos la huella terrestre humana y mapas marinos del impacto humano cumulativo en un análisis global de las presiones antropogénicas que afectan las áreas costeras. De las áreas costeras de todo el mundo, el 15.5% tuvieron una presión antropogénica reducida, principalmente en Canadá, Rusia y Groenlandia. En cambio, el 47.9% de las regiones costeras estuvieron fuertemente afectas por la humanidad, y en la mayoría de los países (84.1%)50% de sus regiones litorales se encuentran degradadas. Casi la mitad (43.3%) de las áreas protegidas en las regiones costeras tienen un grado de exposición a fuertes presiones humanas. Para cumplir los objetivos mundiales de sustentabilidad, todos los países deben emprender mejores acciones para preservar y restaurar las regiones litorales dentro de sus fronteras.

Humans interact with the oceans in diverse and profound ways. The scope, magnitude, footprint and ultimate cumulative impacts of human activities can threaten ocean ecosystems and have changed over time, resulting in new challenges and threats to marine ecosystems. A fundamental gap in understanding how humanity is affecting the oceans is our limited knowledge about the pace of change in cumulative impact on ocean ecosystems from expanding human activities – and the patterns, locations and drivers of most significant change. To help address this, we combined high resolution, annual data on the intensity of 14 human stressors and their impact on 21 marine ecosystems over 11 years (2003–2013) to assess pace of change in cumulative impacts on global oceans, where and how much that pace differs across the ocean, and which stressors and their impacts contribute most to those changes. We found that most of the ocean (59%) is experiencing significantly increasing cumulative impact, in particular due to climate change but also from fishing, land-based pollution and shipping. Nearly all countries saw increases in cumulative impacts in their coastal waters, as did all ecosystems, with coral reefs, seagrasses and mangroves at most risk. Mitigation of stressors most contributing to increases in overall cumulative impacts is urgently needed to sustain healthy oceans.

Sustainability indices are proliferating, both to help synthesize scientific understanding and inform policy. However, it remains poorly understood how such indices are affected by underlying assumptions of the data and modelling approaches used to compute indicator values. Here, we focus on one such indicator, the fisheries goal within the Ocean Health Index (OHI), which evaluates the sustainable provision of food from wild fisheries. We quantify uncertainty in the fisheries goal status arising from the (a) approach for estimating missing data (i.e., fish stocks with no status) and (b) reliance on a data‐limited method (catch‐MSY) to estimate stock status (i.e., B/BMSY). We also compare several other models to estimate B/BMSY, including an ensemble approach, to determine whether alternative models might reduce uncertainty and bias. We find that the current OHI fisheries goal model results in overly optimistic fisheries goal statuses. Uncertainty and bias can be reduced by (a) using a mean (vs. median) gap‐filling approach to estimate missing stock scores and (b) estimating fisheries status using the central tendency from a simulated distribution of status scores generated by a bootstrap approach that incorporates error in B/BMSY. This multitiered approach to measure and describe uncertainty improves the transparency and interpretation of the indicator and allows us to better understand uncertainty around our OHI fisheries model and outputs for country‐level interpretation and use.

Management of the land-sea interface is considered essential for global conservation and sustainability objectives, as coastal regions maintain natural processes that support biodiversity and the livelihood of billions of people. However, assessments of coastal regions have focused on either strictly the terrestrial or marine realm, and as a consequence, we still have a poor understanding of the overall state of Earth’s coastal regions. Here, by integrating the terrestrial human footprint and marine cumulative human impact maps, we provide a global assessment of the anthropogenic pressures affecting coastal areas. Just 15.5% of coastal areas globally can be considered having low anthropogenic pressure, mostly found in Canada, Russia, and Greenland. Conversely, 47.9% of coastal regions are heavily impacted by humanity with most countries (84.1%) having >50% of their coastal regions degraded. Nearly half (43.3%) of protected areas across coastal regions are exposed to high human pressures. In order to meet global sustainability objectives, we identify those nations that must undertake greater actions to preserve and restore coastal regions so as to ensure global sustainable development objectives can be met.

Improving the health of coastal and open sea marine ecosystems represents a substantial challenge for sustainable marine resource management, since it requires balancing human benefits and impacts on the ocean. This challenge is often exacerbated by incomplete knowledge and lack of tools that measure ocean and coastal ecosystem health in a way that allows consistent monitoring of progress towards predefined management targets. The lack of such tools often limits capabilities to enact and enforce effective governance. We introduce the Baltic Health Index (BHI) as a transparent, collaborative and repeatable assessment tool. The Index complements existing, more ecological-oriented, approaches by including a human dimension on the status of the Baltic Sea, an ecosystem impacted by multiple anthropogenic pressures and governed by a multitude of comprehensive national and international policies. Using a large amount of social-ecological data available, we assessed the health of the Baltic Sea for nine goals that represent the status towards set targets, for example, clean waters, biodiversity, food provision, natural products extraction and tourism. Our results indicate that the overall health of the Baltic Sea is suboptimal (a score of 76 out of 100), and a substantial effort is required to reach the management objectives and associated targets. Subregionally, the lowest BHI scores were measured for carbon storage, contaminants and lasting special places (i.e. marine protected areas), albeit with large spatial variation. Overall, the likely future status of all goals in the BHI averaged for the entire Baltic Sea is better than the present status, indicating a positive trend towards a healthier Baltic Sea. However, in some Baltic Sea basins, the trend for specific goals was decreasing, highlighting locations and issues that should be the focus of management priorities. The BHI outcomes can be used to identify both pan-Baltic and subregional scale management priorities and to illustrate the interconnectedness between goals linked by cumulative pressures. Hence, the information provided by the BHI tool and its further development will contribute towards the fulfilment of the UN Agenda 2030 and its Sustainability Development Goals. A free Plain Language Summary can be found within the Supporting Information of this article. A free Plain Language Summary can be found within the Supporting Information of this article.

An ever-growing human footprint Human activities are increasingly affecting the marine environment but understanding how much and in what ways is an extreme challenge given the vastness of this system. O'Hara et al. looked at a suite of human-induced stressors on >1000 marine species over the course of 13 years. They found that species are experiencing increasing levels of stress over more than half of their ranges, with some species having an even higher proportion of their ranges affected. Fishing has the largest impact, but other stressors, such as climate change, are also important and growing. Science , this issue p. 84

Effective management of marine systems requires quantitative tools that can assess the state of the marine social-ecological system and are responsive to management actions and pressures. We applied the Ocean Health Index (OHI) framework to retrospectively assess ocean health in British Columbia annually from 2001 to 2016 for eight goals that represent the values of British Columbia's coastal communities. We found overall ocean health improved over the study period, from 75 (out of 100) in 2001 to 83 in 2016, with scores for inhabited regions ranging from 68 (North Coast, 2002) to 87 (West Vancouver Island, 2011). Highest-scoring goals were Tourism & Recreation (average 94 over the period) and Habitat Services (100); lowest-scoring goals were Sense of Place (61) and Food Provision (64). Significant increases in scores over the time period occurred for Food Provision (+1.7 per year), Sense of Place (+1.4 per year), and Coastal Livelihoods (+0.6 per year), while Habitat Services (-0.01 per year) and Biodiversity (-0.09 per year) showed modest but statistically significant declines. From the results of our time-series analysis, we used the OHI framework to evaluate impacts of a range of management actions. Despite challenges in data availability, we found evidence for the ability of management to reduce pressures on several goals, suggesting the potential of OHI as a tool for assessing the effectiveness of marine resource management to improve ocean health. Our OHI assessment provides an important comprehensive evaluation of ocean health in British Columbia, and our open and transparent process highlights opportunities for improving accessibility of social and ecological data to inform future assessment and management of ocean health.

A mesocosm system was developed to simulate estuarine conditions characteristic of short water-residence time ecosystems of the Pacific Coast of North America, and used to evaluate the response of multiple macrophyte metrics to gradients of NO 3 loading and temperature. Replicated experiments found that few responses could be directly attributed to NO 3 loading up to 6 x ambient. Some response metrics exhibited weak relationships with nutrient loading but could not be resolved with available statistical power. While direct nutrient responses were found for some species-specific metrics (e.g. green macroalgal growth and biomass, tissue N%, etc.), many patterns were confounded with temperature. Temperature generally had a larger effect on response metrics than did nutrient load. Experimental macrophyte communities exhibited community shifts consistent with the predicted effects of nutrient loading at 20 °C, but there was no evidence of other eutrophication symptoms (phytoplankton blooms or hypoxia) due to the short system-residence time. The Z . marina Nutrient Pollution Index (NPI) tracked the NO 3 gradient at 10 °C, but exhibited no response at 20 °C, which may limit the utility of this metric in areas with marked thermal seasonality. Results suggest that teasing apart the influence of temperature and nutrients on the expression of eutrophication symptoms will require complex multi-stressor experiments and the use of indicators that are sensitive across a broad range of conditions.

Coastal marine ecosystems face a host of pressures from both offshore and land-based human activity. Research on terrestrial threats to coastal ecosystems has primarily focused on agricultural runoff, specifically showcasing how fertilizers and livestock waste create coastal eutrophication, harmful algae blooms, or hypoxic or anoxic zones. These impacts not only harm coastal species and ecosystems but also impact human health and economic activities. Few studies have assessed impacts of human wastewater on coastal ecosystems and community health. As such, we lack a comprehensive, fine-resolution, global assessment of human sewage inputs that captures both pathogens and nutrient flows to coastal waters and the potential impacts on coastal ecosystems. To address this gap, we use a new high-resolution geospatial model to measure and map nitrogen (N) and pathogen—fecal indicator organisms (FIO)—inputs from human sewage for ~135,000 watersheds globally. Because solutions depend on the source, we separate nitrogen and pathogen inputs from sewer, septic, and direct inputs. Our model indicates that wastewater adds 6.2Tg nitrogen into coastal waters, which is approximately 40% of total nitrogen from agriculture. Of total wastewater N, 63% (3.9Tg N) comes from sewered systems, 5% (0.3Tg N) from septic, and 32% (2.0Tg N) from direct input. We find that just 25 watersheds contribute nearly half of all wastewater N, but wastewater impacts most coastlines globally, with sewered, septic, and untreated wastewater inputs varying greatly across watersheds and by country. Importantly, model results find that 58% of coral and 88% of seagrass beds are exposed to wastewater N input. Across watersheds, N and FIO inputs are generally correlated. However, our model identifies important fine-grained spatial heterogeneity that highlight potential tradeoffs and synergies essential for management actions. Reducing impacts of nitrogen and pathogens on coastal ecosystems requires a greater focus on where wastewater inputs vary across the planet. Researchers and practitioners can also overlay these global, high resolution, wastewater input maps with maps describing the distribution of habitats and species, including humans, to determine the where the impacts of wastewater pressures are highest. This will help prioritize conservation efforts.Without such information, coastal ecosystems and the human communities that depend on them will remain imperiled.

Summary Carbon offsetting—receiving credit for reducing, avoiding, or sequestering carbon—has become part of the portfolio of solutions to mitigate carbon emissions, and thus climate change, through policy and voluntary markets, primarily by land-based re- or afforestation and preservation [ 1 , 2 ]. However, land is limiting, creating interest in a rapidly growing aquatic farming sector of seaweed aquaculture [ 3 , 4 , 5 ]. Synthesizing data from scientific literature, we assess the extent and cost of scaling seaweed aquaculture to provide sufficient CO2eq sequestration for several climate change mitigation scenarios, with a focus on the food sector—a major source of greenhouse gases [ 6 ]. Given known ecological constraints (nutrients and temperature), we found a substantial suitable area (ca. 48 million km2) for seaweed farming, which is largely unfarmed. Within its own industry, seaweed could create a carbon-neutral aquaculture sector with just 14% (mean = 25%) of current seaweed production (0.001% of suitable area). At a much larger scale, we find seaweed culturing extremely unlikely to offset global agriculture, in part due to production growth and cost constraints. Yet offsetting agriculture appears more feasible at a regional level, especially areas with strong climate policy, such as California (0.065% of suitable area). Importantly, seaweed farming can provide other benefits to coastlines affected by eutrophic, hypoxic, and/or acidic conditions [ 7 , 8 ], creating opportunities for seaweed farming to act as “charismatic carbon” that serves multiple purposes. Seaweed offsetting is not the sole solution to climate change, but it provides an invaluable new tool for a more sustainable future.

People depend on the ocean to provide a range of ecosystem services, including sustaining economies and providing nutrition. We demonstrate how a global ocean health index framework can be applied to a data-limited scenario and modified to incorporate the objectives and context of a developing island nation like Fiji. Although these changes did not have a major effect on the total index value, two goals had substantial changes. The artisanal opportunities goal increased from 46 to 92 as a result of changes to the model for Fiji, which looks at the stock status of artisanally-caught species. The lasting special places sub-goal decreased from 96 to 48, due to the use of Fiji-specific data and reference points that allow policymakers to track progress towards national goals. Fiji scored high for the tourism and recreation goal, but low for the production-oriented natural products goal and mariculture sub-goal, which may reflect national values and development priorities. By measuring ocean health across a portfolio of goals and re-calculating scores over time, we can better understand potential trade-offs between goals. Our approach for measuring ocean health in Fiji highlights pathways for improvements and approaches that may help guide other data-limited countries in assessing ocean health.

Growing international and national focus on quantitatively measuring and improving ocean health has increased the need for comprehensive, scientific, and repeated indicators to track progress towards achieving policy and societal goals. The Ocean Health Index (OHI) is one of the few indicators available for this purpose. Here we present results from five years of annual global assessment for 220 countries and territories, evaluating potential drivers and consequences of changes and presenting lessons learned about the challenges of using composite indicators to measure sustainability goals. Globally scores have shown little change, as would be expected. However, individual countries have seen notable increases or declines due in particular to improvements in the harvest and management of wild-caught fisheries, the creation of marine protected areas (MPAs), and decreases in natural product harvest. Rapid loss of sea ice and the consequent reduction of coastal protection from that sea ice was also responsible for declines in overall ocean health in many Arctic and sub-Arctic countries. The OHI performed reasonably well at predicting near-term future scores for many of the ten goals measured, but data gaps and limitations hindered these predictions for many other goals. Ultimately, all indicators face the substantial challenge of informing policy for progress toward broad goals and objectives with insufficient monitoring and assessment data. If countries and the global community hope to achieve and maintain healthy oceans, we will need to dedicate significant resources to measuring what we are trying to manage.

Reproducibility has long been a tenet of science but has been challenging to achieve-we learned this the hard way when our old approaches proved inadequate to efficiently reproduce our own work. Here we describe how several free software tools have fundamentally upgraded our approach to collaborative research, making our entire workflow more transparent and streamlined. By describing specific tools and how we incrementally began using them for the Ocean Health Index project, we hope to encourage others in the scientific community to do the same-so we can all produce better science in less time.

The Antarctic coast and seas are considered some of the most pristine marine systems on Earth. Their comprehensive assessment is critical because meeting ambitious conservation objectives while maintaining sustainable human uses will be increasingly challenging with growing climate change impacts, recovery from past overharvesting, and potential revision of activities permitted with future revisions of the existing governance structure. We used the Ocean Health Index (OHI) to deliver an integrated assessment of the Antarctic marine ecosystems’ evolving ecological and social dimensions. The OHI provides a framework to evaluate sustainable delivery of benefits people want from healthy oceans by measuring progress towards 10 widely-held societal goals. These goals include, conservation objectives, as well as other objectives, so as to identify tradeoffs across multiple priorities. We adapted the Index to the unique aspects and data availability of Antarctica. OHI scores were calculated for each sub-region defined by the Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR) as well as the region overall. OHI scores for conservation-related goals (biodiversity, clean water) were generally high, though with some stressor impacts (i.e., climate-driven decline of sea-ice, and pathogen pollution). However, a sensitivity test on the sea-ice habitat indicator showed biodiversity scores might be much lower in the vicinity of the Antarctic Peninsula. Preservation of lasting special places, captured in the sense of place sub-goal, scored relatively low due to limited extent of Marine Protected Areas in the Southern Ocean. In several cases, scores are low due to under-utilization of resources, rather than environmentally unsustainable practices (e.g., food provision, natural products, tourism and recreation). However, increased human activities would intensify the risk of pollution, pathogen contamination, and disturbance to wildlife, particularly if compounded with future climate change impacts. Therefore, scores may reflect the need to select more conservative targets for human use, articulated in international treaties, taking future risks into account. Our results highlight the need for more research on both natural and social science aspects of the Antarctic system, as well as the need to evaluate targets under different scenarios, so as to provide robust science-based advice for future decision-making in the region.

Discharge from the ballast tanks of ships is one of the primary vectors of nonindigenous species in marine environments. To mitigate this environmental and economic threat, international, national, and state entities are establishing regulations to limit the concentration of living organisms that may be discharged from the ballast tanks of ships. The proposed discharge standards have ranged from zero detectable organisms to < 10 organisms/ m3. If standard sampling methods are used, verifying whether ballast discharge complies with these stringent standards will be challenging due to the inherent stochasticity of sampling. Furthermore, at low concentrations, very large volumes of water must be sampled to find enough organisms to accurately estimate concentration. Despite these challenges, adequate sampling protocols comprise a critical aspect of establishing standards because they help define the actual risk level associated with a standard. A standard that appears very stringent may be effectively lax if it is paired with an inadequate sampling protocol. We describe some of the statistical issues associated with sampling at low concentrations to help regulators understand the uncertainties of sampling as well as to inform the development of sampling protocols that ensure discharge standards are adequately implemented.

As a vector by which foreign species invade coastal and freshwater waterbodies, ballast water discharge from ships is recognized as a major environmental threat. The International Maritime Organization (IMO) drafted an international treaty establishing ballast water discharge standards based on the number of viable organisms per volume of ballast discharge for different organism size classes. Concerns that the IMO standards are not sufficiently protective have initiated several state and national efforts in the United States to develop more stringent standards. We evaluated seven approaches to establishing discharge standards for the > 50-microm size class: (1) expert opinion/management consensus, (2) zero detectable living organisms, (3) natural invasion rates, (4) reaction-diffusion models, (5) population viability analysis (PVA) models, (6) per capita invasion probabilities (PCIP), and (7) experimental studies. Because of the difficulty in synthesizing scientific knowledge in an unbiased and transparent fashion, we recommend the use of quantitative models instead of expert opinion. The actual organism concentration associated with a "zero detectable organisms" standard is defined by the statistical rigor of its monitoring program; thus it is not clear whether such a standard is as stringent as other standards. For several reasons, the natural invasion rate, reaction-diffusion, and experimental approaches are not considered suitable for generating discharge standards. PVA models can be used to predict the likelihood of establishment of introduced species but are limited by a lack of population vital rates for species characteristic of ballast water discharges. Until such rates become available, PVA models are better suited to evaluate relative efficiency of proposed standards rather than predicting probabilities of invasion. The PCIP approach, which is based on historical invasion rates at a regional scale, appears to circumvent many of the indicated problems, although it may underestimate invasions by asexual and parthenogenic species. Further research is needed to better define propagule dose-responses, densities at which Allee effects occur, approaches to predicting the likelihood of invasion from multi-species introductions, and generation of formal comparisons of approaches using standardized scenarios.

To reduce ballast water-borne aquatic invasions worldwide, the International Maritime Organization and United States Coast Guard have each proposed discharge standards specifying maximum concentrations of living biota that may be released in ships’ ballast water (BW), but these regulations still lack guidance for standardized type approval and compliance testing of treatment systems. Verifying whether BW meets a discharge standard poses significant challenges. Properly treated BW will contain extremely sparse numbers of live organisms, and robust estimates of rare events require extensive sampling efforts. A balance of analytical rigor and practicality is essential to determine the volume of BW that can be reasonably sampled and processed, yet yield accurate live counts. We applied statistical modeling to a range of sample volumes, plankton concentrations, and regulatory scenarios (i.e., levels of type I and type II errors), and calculated the statistical power of each combination to detect noncompliant discharge concentrations. The model expressly addresses the roles of sampling error, BW volume, and burden of proof on the detection of noncompliant discharges in order to establish a rigorous lower limit of sampling volume. The potential effects of recovery errors (i.e., incomplete recovery and detection of live biota) in relation to sample volume are also discussed.

Human pressures on the ocean are thought to be increasing globally, yet we know little about their patterns of cumulative change, which pressures are most responsible for change, and which places are experiencing the greatest increases. Managers and policymakers require such information to make strategic decisions and monitor progress towards management objectives. Here we calculate and map recent change over 5 years in cumulative impacts to marine ecosystems globally from fishing, climate change, and ocean- and land-based stressors. Nearly 66% of the ocean and 77% of national jurisdictions show increased human impact, driven mostly by climate change pressures. Five percent of the ocean is heavily impacted with increasing pressures, requiring management attention. Ten percent has very low impact with decreasing pressures. Our results provide large-scale guidance about where to prioritize management efforts and affirm the importance of addressing climate change to maintain and improve the condition of marine ecosystems., Human pressure on the ocean is thought to be increasing globally, yet the magnitude and patterns of these changes are largely unknown. Here, the authors produce a global map of change in cumulative human pressures over the past 5 years, and show that ∼66% of the ocean has experienced elevated human impact.

Oxygen is critically important for catabolic ATP generation but is also a dangerous source of reactive oxygen species. Insects respond to short-term exposure to hypoxia or hyperoxia with compensatory changes in spiracular opening and ventilation that reduce variation in internal Po2. Below critical Po2 values (Pc), nitric oxide and hypoxia inducible factor (HIF)-mediated pathways induce long-term responses such as compensatory tracheal growth, suppressed development, and acclimation of ventilation. Pc values are strongly affected by activity and ontogeny, due to changes in the ratio of tracheal conductance to metabolic rate. Although growth rates and development are suppressed by significant hypoxia in all species studied to date, adult body size is only affected in some species. Severe hyperoxia causes major oxidative stress and reduces survival, while moderate hyperoxia increases development times and body sizes in some species by unknown mechanisms.

Synopsis Numerous physical parameters that influence insect physiology vary substantially with altitude, including temperature, air density, and oxygen partial pressure. Here, we review existing literature and present new empirical data to better characterize the high-altitude environment, and then consider how this environment affects the physiology and evolution of insects. Using weather balloon data from fifty-three sites across the globe, we estimate a mean altitudinal temperature lapse rate of 6.0 � C/km. We also present empirically determined lapse rates for PO2 and air density. The temperature decline with elevation may substantially compromise insect thermoregulation at high altitude. However, heat-transfer models predict that lower air density at elevation reduces convective heat loss of insects by to a surprisingly large degree. This effect combined with behavioral thermoregulation and the availability of buffered microhabitats make the net thermal consequences of highaltitude residence strongly context-specific. The decline in PO2 with elevation may compromise insect development and physiology, but its effects are difficult to predict without simultaneously considering temperature and air density. Flying insects compensate for low air densities with both short-term responses, such as increased stroke amplitude (but not wingbeat frequency), and with long-term developmental and/or evolutionary increases in wing size relative to body size. Finally, in contrast to predictions based on Bergmann’s Rule, a literature survey of thirty-six insect species suggests that those living in colder, higher altitudes do not tend to have larger body sizes. Rapid changes in the physical environment can profoundly alter biological communities across altitudinal gradients. For example, mean temperature drops by approximately 24 � C from the base of the Sierra Nevadas (200 m) to the peak of Mount Dana (3979 m) in Yosemite National Park, California. The altitudinal temperature change associated with this short horizontally projected distance (97 km) is roughly equivalent to traveling over 4500 km in latitude (Hopkins, 1938). This rapid temperature reduction is likely a primary factor underlying the striking altitudinal changes in plant and animal communities. However, other physical factors such as oxygen, air density, humidity, and solar radiation may also shape high-altitude insect communities, but have been largely ignored. In this review, we discuss the rapid changes in the physical factors that characterize high altitude, and explore how these gradients influence the physiology and evolution of arthropods, and particularly of insects, living at high altitude. Rather than exhaustively review topics for which data are abundant, we primarily present hypotheses for which data are scarce or nonexistent in the hopes of encouraging future research.

A habitat-based framework is a practical method for developing models (or, ecological production functions, EPFs) to describe the spatial distribution of ecosystem services. To generate EPFs for Yaquina estuary, Oregon, USA, we compared bird use patterns among intertidal habitats. Visual censuses were used to quantify abundance of bird groups and general species richness in: Zostera marina (eelgrass), Upogebia (mud shrimp)/mudflat, Neotrypaea (ghost shrimp)/sandflat, Zostera japonica (Japanese eelgrass), and low marsh estuarine habitats. Also assessed were (1) spatial variation within a habitat along the estuary gradient and, (2) temporal variation based on bi-monthly samples over a year at five tidal ranges. Z. marina was an important estuarine habitat based on nearly all metrics of bird use, except for shorebird densities. This suggests that reductions in native eelgrass habitat may reduce the abundance and diversity of birds in Yaquina estuary. Our results suggest that a habitat based assessment approach is generally feasible for developing relative EPFs related to the presence of birds within estuarine systems.

Although higher temperatures strongly stimulate ectothermic metabolic rates, they only slightly increase oxygen diffusion rates and decrease oxygen solubility. Consequently, we predicted that insect gas exchange systems would have more difficulty meeting tissue oxygen demands at higher temperatures. In this study, Drosophila melanogaster were reared from egg to adult in hyperoxic (40%), hypoxic (10%), and normoxic (21%) conditions and in temperatures ranging from 15 degrees -31.5 degrees C to examine the interactive effect of temperature and oxygen on development. Hyperoxia generally increased mass and growth rate at higher rearing temperatures. At lower rearing temperatures, however, hyperoxia had a very small effect on mass, did not affect growth rate, and lengthened time to eclosion. Relative to normoxia, flies reared in hypoxic conditions were generally smaller (mass and thorax length), had longer eclosion times, slower growth rates, and reduced survival. At cooler temperatures, hypoxia had relatively modest or nonsignificant effects on development, while at higher temperatures, the effects of hypoxia were large. These results suggest that higher temperatures reduce oxygen delivery capacity relative to tissue oxygen needs, which may partially explain why ectotherms are smaller when development occurs at higher temperatures.

Despite decades of research on the evolution of thermal physiology, at least one fundamental issue remains unresolved: whether the maximal performance of a genotype depends on its optimal temperature. One school argues that warm‐adapted genotypes will outperform cold‐adapted genotypes because high temperatures inevitably accelerate chemical reactions. Yet another school holds that biochemical adaptation can compensate for thermodynamic effects on performance. Here, we briefly discuss this theoretical debate and then summarize empirical studies that address whether hotter is better. In general, comparative and experimental studies support the view that hotter is better. Furthermore, recent modeling has shown that thermodynamic constraints impose unique selective pressures on thermal sensitivity. Nevertheless, the thermodynamic effect on maximal performance varies greatly among traits and taxa, suggesting the need to develop a more sophisticated view of thermodynamic constraints.

We use a factorial experimental design to test whether rearing at colder temperatures shifts the lower thermal envelope for flight of Drosophila melanogaster Meigen to colder temperatures. D. melanogaster that developed in colder temperatures (15 degrees C) had a significant flight advantage in cold air compared to flies that developed in warmer temperatures (28 degrees C). At 14 degrees C, cold-reared flies failed to perform a take-off flight approximately 47% of the time whereas warm-reared flies failed approximately 94% of the time. At 18 degrees C, cold- and warm-reared flies performed equally well. We also compared several traits in cold- and warm-developing flies to determine if cold-developing flies had better flight performance at cold temperatures due to changes in body mass, wing length, wing loading, relative flight muscle mass or wing-beat frequency. The improved ability to fly at low temperatures was associated with a dramatic increase in wing area and an increase in wing length (after controlling for wing area). Flies that developed at 15 degrees C had approximately 25% more wing area than similarly sized flies that developed at 28 degrees C. Cold-reared flies had slower wing-beat frequencies than similarly sized flies from warmer developmental environments, whereas other traits did not vary with developmental temperature. These results demonstrate that developmental plasticity in wing dimensions contributes to the improved flight performance of D. melanogaster at cold temperatures, and ultimately, may help D. melanogaster live in a wide range of thermal environments.

International and regional policies aimed at managing ocean ecosystem health need quantitative and comprehensive indices to synthesize information from a variety of sources, consistently measure progress, and communicate with key constituencies and the public. Here we present the second annual global assessment of the Ocean Health Index, reporting current scores and annual changes since 2012, recalculated using updated methods and data based on the best available science, for 221 coastal countries and territories. The Index measures performance of ten societal goals for healthy oceans on a quantitative scale of increasing health from 0 to 100, and combines these scores into a single Index score, for each country and globally. The global Index score improved one point (from 67 to 68), while many country-level Index and goal scores had larger changes. Per-country Index scores ranged from 41–95 and, on average, improved by 0.06 points (range -8 to +12). Globally, average scores increased for individual goals by as much as 6.5 points (coastal economies) and decreased by as much as 1.2 points (natural products). Annual updates of the Index, even when not all input data have been updated, provide valuable information to scientists, policy makers, and resource managers because patterns and trends can emerge from the data that have been updated. Changes of even a few points indicate potential successes (when scores increase) that merit recognition, or concerns (when scores decrease) that may require mitigative action, with changes of more than 10–20 points representing large shifts that deserve greater attention. Goal scores showed remarkably little covariance across regions, indicating low redundancy in the Index, such that each goal delivers information about a different facet of ocean health. Together these scores provide a snapshot of global ocean health and suggest where countries have made progress and where a need for further improvement exists.

Diverse biochemical and physiological adaptations enable different species of ectotherms to survive and reproduce in very different temperature regimes, but whether these adaptations fully compensate for the thermodynamically depressing effects of low temperature on rates of biological processes is debated. If such adaptations are fully compensatory, then temperature-dependent processes (e.g., digestion rate, population growth rate) of cold-adapted species will match those of warm-adapted species when each is measured at its own optimal temperature. Here we show that cold-adapted insect species have much lower maximum rates of population growth than do warm-adapted species, even when we control for phylogenetic relatedness. This pattern also holds when we use a structural-equation model to analyze alternative hypotheses that might otherwise explain this correlation. Thus, although physiological adaptations enable some insects to survive and reproduce at low temperatures, these adaptations do not overcome the "tyranny" of thermodynamics, at least for rates of population increase. Indeed, the sensitivity of population growth rates of insects to temperature is even greater than predicted by a recent thermodynamic model. Our findings suggest that adaptation to temperature inevitably alters the population dynamics of insects. This result has broad evolutionary and ecological consequences.

SUMMARYThe alpine environment is likely to challenge insect locomotion because of low mean temperatures and reduced barometric pressure. In this study, we measured the direct and interactive effects of these factors on walking and flight performance of wild-caught Drosophila melanogaster Meigen. We found that decreased temperature and decreased air pressure both reduced walking speed and flight performance. Flies walked more slowly at 18°C and in the lowest air pressure treatment (34 kPa). This treatment, equivalent in air pressure to the top of Mount Everest, was the only air pressure that significantly reduced fly walking speed. Therefore, walking performance in the wild is likely limited by temperature, but not oxygen availability. In contrast to walking performance, low but ecologically realistic air pressures dramatically reduced overall flight performance. The effects of reduced air pressure on flight performance were more pronounced at colder temperatures. Reduced flight performance in high altitude conditions was primarily driven by an increased reluctance for flies to initiate flight rather than outright failure to fly. Such reluctance to fly in high altitude conditions may in part explain the prevalence of aptery and brachyptery in high altitude insects. The observed interactive effects of temperature and air pressure on flight performance confirm the importance of simultaneously manipulating both of these factors when studying the impact of altitudinal conditions on insect physiology and behavior.

Development time is a critical life-history trait that has profound effects on organism fitness and on population growth rates. For ectotherms, development time is strongly influenced by temperature and is predicted to scale with body mass to the quarter power based on 1) the ontogenetic growth model of the metabolic theory of ecology which describes a bioenergetic balance between tissue maintenance and growth given the scaling relationship between metabolism and body size, and 2) numerous studies, primarily of vertebrate endotherms, that largely support this prediction. However, few studies have investigated the allometry of development time among invertebrates, including insects. Abundant data on development of diverse insects provides an ideal opportunity to better understand the scaling of development time in this ecologically and economically important group. Insects develop more quickly at warmer temperatures until reaching a minimum development time at some optimal temperature, after which development slows. We evaluated the allometry of insect development time by compiling estimates of minimum development time and optimal developmental temperature for 361 insect species from 16 orders with body mass varying over nearly 6 orders of magnitude. Allometric scaling exponents varied with the statistical approach: standardized major axis regression supported the predicted quarter-power scaling relationship, but ordinary and phylogenetic generalized least squares did not. Regardless of the statistical approach, body size alone explained less than 28% of the variation in development time. Models that also included optimal temperature explained over 50% of the variation in development time. Warm-adapted insects developed more quickly, regardless of body size, supporting the "hotter is better" hypothesis that posits that ectotherms have a limited ability to evolutionarily compensate for the depressing effects of low temperatures on rates of biological processes. The remaining unexplained variation in development time likely reflects additional ecological and evolutionary differences among insect species.

Despite the potential for diet to affect organismal acid-base status, especially in herbivores, little is known about the effects of diet on acid-base loading and excretion. We tested the effects of diet on acid-base loading and excretion in grasshoppers by (a) comparing the fecal acid-base content of 15 grasshopper species collected from the field and (b) comparing fecal acid-base excretion rates of Schistocerca americana grasshoppers fed vegetable diets that differed in their ashed and raw acid-base contents. The field experiments indicated that grass-feeding species excrete fairly neutral fecal pellets, while forb/mixed-feeding species vary widely in their fecal acid-base contents. In the laboratory experiment, acid-base excretion rates were positively correlated with dietary ashed base intake rates but were not correlated with the acid-base content of raw, unashed diet or feeding rate. These experiments suggest that some diets could strongly challenge the acid-base homeostasis of herbivores; in some grasshoppers, dietary acid-base loads could produce certainly lethal 1-unit changes in average body pH within 6 h if they were not excreted.