Going with the flow: the role of ocean circulation in global marine ecosystems under a changing climate

Ocean warming, acidification, deoxygenation and reduced productivity are widely considered to be the major stressors to ocean ecosystems induced by emissions of CO₂. However, an overlooked stressor is the change in ocean circulation in response to climate change. Strong changes in the intensity and position of the western boundary currents have already been observed, and the consequences of such changes for ecosystems are beginning to emerge. In this study, we address climatically induced changes in ocean circulation on a global scale but relevant to propagule dispersal for species inhabiting global shelf ecosystems, using a high‐resolution global ocean model run under the IPCC RCP 8.5 scenario. The ¼ degree model resolution allows improved regional realism of the ocean circulation beyond that of available CMIP5‐class models. We use a Lagrangian approach forced by modelled ocean circulation to simulate the circulation pathways that disperse planktonic life stages. Based on trajectory backtracking, we identify present‐day coastal retention, dominant flow and dispersal range for coastal regions at the global scale. Projecting into the future, we identify areas of the strongest projected circulation change and present regional examples with the most significant modifications in their dominant pathways. Climatically induced changes in ocean circulation should be considered as an additional stressor of marine ecosystems in a similar way to ocean warming or acidification.     

全球变化对海洋生态系统初级生产关键过程的影响

工业革命以来, 不断加剧的人类活动所引起的大气CO₂浓度增加、温度上升等全球变化问题, 正使得海洋生态系统面临着前所未有的压力。该文通过文献计量的方法分析了国内外的研究现状, 简要地回顾了全球变化对海洋生态系统影响研究的发展简史, 并聚焦海洋暖化、海洋酸化和富营养化与缺氧这三个核心研究方向, 重点阐述了它们对海洋生态系统初级生产的关键过程的影响, 总结了已取得的重要进展以及存在的主要问题, 最后提出前沿展望。     

Effects of global change on key processes of primary production in marine ecosystems

工业革命以来, 不断加剧的人类活动所引起的大气CO₂浓度增加、温度上升等全球变化问题, 正使得海洋生态系统面临着前所未有的压力。该文通过文献计量的方法分析了国内外的研究现状, 简要地回顾了全球变化对海洋生态系统影响研究的发展简史, 并聚焦海洋暖化、海洋酸化和富营养化与缺氧这三个核心研究方向, 重点阐述了它们对海洋生态系统初级生产的关键过程的影响, 总结了已取得的重要进展以及存在的主要问题, 最后提出前沿展望。     

Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes

Analysis of Permian–Triassic brachiopod diversity and body size changes from different water depths spanning the continental shelf to basinal facies in South China provides insights into the process of environmental deterioration. Comparison of the temporal changes of brachiopod diversity between deepwater and shallow‐water facies demonstrates that deepwater brachiopods disappeared earlier than shallow‐water brachiopods. This indicates that high environmental stress commenced first in deepwater settings and later extended to shallow waters. This environmental stress is attributed to major volcanic eruptions, which first led to formation of a stratified ocean and a chemocline in the outer shelf and deeper water environments, causing the disappearance of deep marine benthos including brachiopods. The chemocline then rapidly migrated upward and extended to shallow waters, causing widespread mass extinction of shallow marine benthos. We predict that the spatial and temporal patterns of earlier onset of disappearance/extinction and ecological crisis in deeper water ecosystems will be recorded during other episodes of rapid global warming.     

Organism activity levels predict marine invertebrate survival during ancient global change extinctions

Multistressor global change, the combined influence of ocean warming, acidification, and deoxygenation, poses a serious threat to marine organisms. Experimental studies imply that organisms with higher levels of activity should be more resilient, but testing this prediction and understanding organism vulnerability at a global scale, over evolutionary timescales, and in natural ecosystems remain challenging. The fossil record, which contains multiple extinctions triggered by multistressor global change, is ideally suited for testing hypotheses at broad geographic, taxonomic, and temporal scales. Here, I assess the importance of activity level for survival of well‐skeletonized benthic marine invertebrates over a 100‐million‐year‐long interval (Permian to Jurassic periods) containing four global change extinctions, including the end‐Permian and end‐Triassic mass extinctions. More active organisms, based on a semiquantitative score incorporating feeding and motility, were significantly more likely to survive during three of the four extinction events (Guadalupian, end‐Permian, and end‐Triassic). In contrast, activity was not an important control on survival during nonextinction intervals. Both the end‐Permian and end‐Triassic mass extinctions also triggered abrupt shifts to increased dominance by more active organisms. Although mean activity gradually returned toward pre‐extinction values, the net result was a permanent ratcheting of ecosystem‐wide activity to higher levels. Selectivity patterns during ancient global change extinctions confirm the hypothesis that higher activity, a proxy for respiratory physiology, is a fundamental control on survival, although the roles of specific physiological traits (such as extracellular pCO₂ or aerobic scope) cannot be distinguished. Modern marine ecosystems are dominated by more active organisms, in part because of selectivity ratcheting during these ancient extinctions, so on average may be less vulnerable to global change stressors than ancient counterparts. However, ancient extinctions demonstrate that even active organisms can suffer major extinction when the intensity of environmental disruption is intense.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

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.     

Mass extinctions and ocean acidification: biological constraints on geological dilemmas

The five mass extinction events that the earth has so far experienced have impacted coral reefs as much or more than any other major ecosystem. Each has left the Earth without living reefs for at least four million years, intervals so great that they are commonly referred to as ‘reef gaps’ (geological intervals where there are no remnants of what might have been living reefs). The causes attributed to each mass extinction are reviewed and summarised. When these causes and the reef gaps that follow them are examined in the light of the biology of extant corals and their Pleistocene history, most can be discarded. Causes are divided into (1) those which are independent of the carbon cycle: direct physical destruction from bolides, ‘nuclear winters’ induced by dust clouds, sea-level changes, loss of area during sea-level regressions, loss of biodiversity, low and high temperatures, salinity, diseases and toxins and extraterrestrial events and (2) those linked to the carbon cycle: acid rain, hydrogen sulphide, oxygen and anoxia, methane, carbon dioxide, changes in ocean chemistry and pH. By process of elimination, primary causes of mass extinctions are linked in various ways to the carbon cycle in general and ocean chemistry in particular with clear association with atmospheric carbon dioxide levels. The prospect of ocean acidification is potentially the most serious of all predicted outcomes of anthropogenic carbon dioxide increase. This study concludes that acidification has the potential to trigger a sixth mass extinction event and to do so independently of anthropogenic extinctions that are currently taking place.     

The future of the northeast Atlantic benthic flora in a high CO2 world

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO₂ emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf‐forming seaweeds.     

全球气候变化下的海洋浮游植物多样性

理解全球气候变化对地球生态系统的影响是全世界广泛关注的问题, 而相比于陆地生态系统, 海洋生态系统对全球气候变化更为敏感。全球气候变化对海洋的影响主要表现在海洋暖化、海洋酸化、大洋环流系统的改变、海平面上升、紫外线辐射增强等方面。浮游植物是海洋生态系统最重要的初级生产者, 同时对海洋碳循环起到举足轻重的作用, 其对全球气候变化的响应主要体现在物种分布、初级生产力、群落演替、生物气候学等方面。具体表现在以下方面: 暖水种的分布范围在扩大, 冷水种分布范围在缩小; 浮游植物全球初级生产力降低; 浮游植物群落会向细胞体积更小的物种占优势的方向转变; 浮游植物水华发生的时间提前、强度增强; 一些有害物种水华的发生频率也会增加; 海洋表层海水的酸化会影响浮游植物特别是钙化类群的生长和群落多样性; 紫外辐射增强对浮游植物的生长起到抑制作用; 厄尔尼诺、拉尼娜、降水量的增加通常抑制浮游植物生长。浮游植物生长和分布的变化会体现在多样性的各个层面上。对于浮游植物在全球变化各种驱动因子下的生理生态学和长周期变动观测等是今后研究的重要方向, 也将为理解全球变化下的浮游植物-多样性-生态系统响应与反馈机制提供基本信息。     

The reef-building coral Siderastrea siderea exhibits parabolic responses to ocean acidification and warming

Anthropogenic increases in atmospheric CO₂ over this century are predicted to cause global average surface ocean pH to decline by 0.1–0.3 pH units and sea surface temperature to increase by 1–4°C. We conducted controlled laboratory experiments to investigate the impacts of CO₂-induced ocean acidification (pCO₂ = 324, 477, 604, 2553 µatm) and warming (25, 28, 32°C) on the calcification rate of the zooxanthellate scleractinian coral Siderastrea siderea, a widespread, abundant and keystone reef-builder in the Caribbean Sea. We show that both acidification and warming cause a parabolic response in the calcification rate within this coral species. Moderate increases in pCO₂ and warming, relative to near-present-day values, enhanced coral calcification, with calcification rates declining under the highest pCO₂ and thermal conditions. Equivalent responses to acidification and warming were exhibited by colonies across reef zones and the parabolic nature of the corals'' response to these stressors was evident across all three of the experiment''s 30-day observational intervals. Furthermore, the warming projected by the Intergovernmental Panel on Climate Change for the end of the twenty-first century caused a fivefold decrease in the rate of coral calcification, while the acidification projected for the same interval had no statistically significant impact on the calcification rate—suggesting that ocean warming poses a more immediate threat than acidification for this important coral species.     

Defining CO2 and O2 syndromes of marine biomes in the Anthropocene

Research efforts have intensified to foresee the prospects for marine biomes under climate change and anthropogenic drivers over varying temporal and spatial scales. Parallel with these efforts is the utilization of terminology, such as ‘ocean acidification’ (OA) and ‘ocean deoxygenation’ (OD), that can foster rapid comprehension of complex processes driving carbon dioxide (CO₂) and oxygen (O₂) concentrations in the global ocean and thus, are now widely used in discussions within and beyond academia. However, common usage of the terms ‘acidification’ and ‘deoxygenation’ alone are subjective and, without adequate contextualization, have the potential to mislead inferences over drivers that may ultimately shape the future state of marine ecosystems. Here we clarify the usage of the terms OA and OD as global, climate change‐driven processes and discuss the various attributes of elevated CO₂ and reduced O₂ syndromes common to coastal ecosystems. We support the use of the existing terms ‘coastal acidification’ and ‘coastal deoxygenation’ because they help differentiate the sometimes rapid and extreme nature of CO₂ and O₂ syndromes in coastal ecosystems from the global, climate change‐driven processes of OA and OD. Given the complexity and breadth of the processes involved in altering CO₂ and O₂ concentrations across marine ecosystems, we provide a workflow to enable contextualization and clarification of the usage of existing terms and highlight the close link between these two gases across spatial and temporal scales in the ocean. These distinctions are crucial to guide effective communication of research within the scientific community and guide policymakers responsible for intervening on the drivers to secure desirable future ocean states.     

Environmental DNA metabarcoding reveals winners and losers of global change in coastal waters

Studies of the ecological effects of global change often focus on one or a few species at a time. Consequently, we know relatively little about the changes underway at real-world scales of biological communities, which typically have hundreds or thousands of interacting species. Here, we use COI mtDNA amplicons from monthly samples of environmental DNA to survey 221 planktonic taxa along a gradient of temperature, salinity, dissolved oxygen and carbonate chemistry in nearshore marine habitat. The result is a high-resolution picture of changes in ecological communities using a technique replicable across a wide variety of ecosystems. We estimate community-level differences associated with time, space and environmental variables, and use these results to forecast near-term community changes due to warming and ocean acidification. We find distinct communities in warmer and more acidified conditions, with overall reduced richness in diatom assemblages and increased richness in dinoflagellates. Individual taxa finding more suitable habitat in near-future waters are more taxonomically varied and include the ubiquitous coccolithophore Emiliania huxleyi and the harmful dinoflagellate Alexandrium sp. These results suggest foundational changes for nearshore food webs under near-future conditions.     

Generality in multispecies responses to ocean acidification revealed through multiple hypothesis testing

Decades of research have demonstrated that many calcifying species are negatively affected by ocean acidification, a major anthropogenic threat in marine ecosystems. However, even closely related species may exhibit different responses to ocean acidification and less is known about the drivers that shape such variation in different species. Here, we examine the drivers of physiological performance under ocean acidification in a group of five species of turf‐forming coralline algae. Specifically, quantitating the relative weight of evidence for each of ten hypotheses, we show that variation in coralline calcification and photosynthesis was best explained by allometric traits. Across ocean acidification conditions, larger individuals (measured as noncalcified mass) had higher net calcification and photosynthesis rates. Importantly, our approach was able to not only identify the aspect of size that drove the performance of coralline algae, but also determined that responses to ocean acidification were not dependent on species identity, evolutionary relatedness, habitat, shape, or structural composition. In fact, we found that failure to test multiple, alternative hypotheses would underestimate the generality of physiological performances, leading to the conclusion that each species had different baseline performance under ocean acidification. Testing among alternative hypotheses is an essential step toward determining the generalizability of experiments across taxa and identifying common drivers of species responses to global change.     

Early-life exposure to climate change impairs tropical shark survival

Sharks are one of the most threatened groups of marine animals worldwide, mostly owing to overfishing and habitat degradation/loss. Although these cartilaginous fish have evolved to fill many ecological niches across a wide range of habitats, they have limited capability to rapidly adapt to human-induced changes in their environments. Contrary to global warming, ocean acidification was not considered as a direct climate-related threat to sharks. Here we show, for the first time, that an early ontogenetic acclimation process of a tropical shark (Chiloscyllium punctatum) to the projected scenarios of ocean acidification (ΔpH = 0.5) and warming (+4°C; 30°C) for 2100 elicited significant impairments on juvenile shark condition and survival. The mortality of shark embryos at the present-day thermal scenarios was 0% both at normocapnic and hypercapnic conditions. Yet routine metabolic rates (RMRs) were significantly affected by temperature, pH and embryonic stage. Immediately after hatching, the Fulton condition of juvenile bamboo sharks was significantly different in individuals that experienced future warming and hypercapnia; 30 days after hatching, survival rapidly declined in individuals experiencing both ocean warming and acidification (up to 44%). The RMR of juvenile sharks was also significantly affected by temperature and pH. The impact of low pH on ventilation rates was significant only under the higher thermal scenario. This study highlights the need of experimental-based risk assessments of sharks to climate change. In other words, it is critical to directly assess risk and vulnerability of sharks to ocean acidification and warming, and such effort can ultimately help managers and policy-makers to take proactive measures targeting most endangered species.     

Could some coral reefs become sponge reefs as our climate changes?

Coral reefs across the world have been seriously degraded and have a bleak future in response to predicted global warming and ocean acidification (OA). However, this is not the first time that biocalcifying organisms, including corals, have faced the threat of extinction. The end‐Triassic mass extinction (200 million years ago) was the most severe biotic crisis experienced by modern marine invertebrates, which selected against biocalcifiers; this was followed by the proliferation of another invertebrate group, sponges. The duration of this sponge‐dominated period far surpasses that of alternative stable‐ecosystem or phase‐shift states reported on modern day coral reefs and, as such, a shift to sponge‐dominated reefs warrants serious consideration as one future trajectory of coral reefs. We hypothesise that some coral reefs of today may become sponge reefs in the future, as sponges and corals respond differently to changing ocean chemistry and environmental conditions. To support this hypothesis, we discuss: (i) the presence of sponge reefs in the geological record; (ii) reported shifts from coral‐ to sponge‐dominated systems; and (iii) direct and indirect responses of the sponge holobiont and its constituent parts (host and symbionts) to changes in temperature and pH. Based on this evidence, we propose that sponges may be one group to benefit from projected climate change and ocean acidification scenarios, and that increased sponge abundance represents a possible future trajectory for some coral reefs, which would have important implications for overall reef functioning.     

Human-induced salinity changes impact marine organisms and ecosystems

Climate change is fundamentally altering marine and coastal ecosystems on a global scale. While the effects of ocean warming and acidification on ecology and ecosystem functions and services are being comprehensively researched, less attention is directed toward understanding the impacts of human-driven ocean salinity changes. The global water cycle operates through water fluxes expressed as precipitation, evaporation, and freshwater runoff from land. Changes to these in turn modulate ocean salinity and shape the marine and coastal environment by affecting ocean currents, stratification, oxygen saturation, and sea level rise. Besides the direct impact on ocean physical processes, salinity changes impact ocean biological functions with the ecophysiological consequences are being poorly understood. This is surprising as salinity changes may impact diversity, ecosystem and habitat structure loss, and community shifts including trophic cascades. Climate model future projections (of end of the century salinity changes) indicate magnitudes that lead to modification of open ocean plankton community structure and habitat suitability of coral reef communities. Such salinity changes are also capable of affecting the diversity and metabolic capacity of coastal microorganisms and impairing the photosynthetic capacity of (coastal and open ocean) phytoplankton, macroalgae, and seagrass, with downstream ramifications on global biogeochemical cycling. The scarcity of comprehensive salinity data in dynamic coastal regions warrants additional attention. Such datasets are crucial to quantify salinity-based ecosystem function relationships and project such changes that ultimately link into carbon sequestration and freshwater as well as food availability to human populations around the globe. It is critical to integrate vigorous high-quality salinity data with interacting key environmental parameters (e.g., temperature, nutrients, oxygen) for a comprehensive understanding of anthropogenically induced marine changes and its impact on human health and the global economy.     

Silurian carbonate platforms and extinction events—ecosystem changes exemplified from Gotland, Sweden

Recent and ancient carbonate platforms are major marine ecosystems, built by various carbonate-secreting organisms with different sensitivity for environmental change. For this reason, carbonate platforms are excellent sensors for changes in contemporaneous marine environments. A variety of ecosystem changes in carbonate platforms have previously been recognised in the aftermath of mass extinction events. This paper addresses how two Silurian extinction events among graptolites, conodonts, and pentamerid brachiopods can be related to changes in the style of carbonate production and general evolution of low latitude carbonate platforms in a similar way as previously reported from the major five mass extinctions of the Phanerozoic. Strata formed on Gotland during the Mulde and Lau events share remarkably many similarities but are strikingly different in composition compared to other strata on the island. The event-related strata is characterised by the sudden appearance of widespread oolites, deviating reef composition, flat-pebble conglomerates, abundant micro- and macro-oncoids, stromatolites, and other microbial facies suggesting decreased bioturbation levels in contemporaneous shelf seas. Importantly, these changes can be tied to high-resolution biostratigraphic frameworks and global stable isotope excursions. The anomalous intervals may therefore be searched for elsewhere in order to test their regional or global significance.     

Temporal effects of ocean warming and acidification on coral–algal competition

Coral Reefs - While there is an ever-expanding list of impacts on coral reefs as a result of ocean warming and acidification, there is little information on how these global changes influence...