411

Ocean Systems

Coordinating Lead Authors:

Hans-O. Pörtner (Germany), David M. Karl (USA)

Lead Authors:

Philip W. Boyd (New Zealand/Australia), William W.L. Cheung (Canada),

Salvador E. Lluch-Cota (Mexico), Yukihiro Nojiri (Japan), Daniela N. Schmidt (UK),

Peter O. Zavialov (Russian Federation)

Contributing Authors:

Jürgen Alheit (Germany), Javier Aristegui (Spain), Claire Armstrong (Norway),

Gregory Beaugrand (France), Vsevolod Belkovich (Russian Federation), Chris Bowler (France),

Peter Brewer (USA), Matthew Church (USA), Sarah R. Cooley (USA), Pablo del Monte-Luna

(Mexico), Martin Edwards (UK), Mikhail Flint (Russian Federation), Michael J. Follows (USA),

Thomas Frölicher (Switzerland), Elizabeth A. Fulton (Australia), Jean-Pierre Gattuso (France),

Ove Hoegh-Guldberg (Australia), Eileen E. Hofmann (USA), Andrew H. Knoll (USA),

Lisa A. Levin (USA), Lena Menzel (Germany), Coleen L. Moloney (South Africa), R. Ian Perry

(Canada), Elvira S. Poloczanska (Australia), J. Murray Roberts (UK), Björn Rost (Germany),

Jorge L. Sarmiento (USA), Jan Sedláček (Switzerland), Daniela Storch (Germany),

Christian Wiencke (Germany), Astrid C. Wittmann (Germany)

Review Editors:

Kenneth F. Drinkwater (Norway), Alexander Polonsky (Ukraine)

Volunteer Chapter Scientists:

Lena Menzel (Germany), Astrid C. Wittmann (Germany)

This chapter should be cited as:

Pörtner

, H.-O., D.M. Karl, P.W. Boyd, W.W.L. Cheung, S.E. Lluch-Cota, Y. Nojiri, D.N. Schmidt, and P.O. Zavialov,

2014: Ocean systems. In: Climate Change 2014: Impacts, Adaptation, and Vulnerability. Part A: Global and

Sectoral Aspects. Contribution of Working Group II to the Fifth Assessment Report of the Intergovernmental

Panel on Climate Change [Field, C.B., V.R. Barros, D.J. Dokken, K.J. Mach, M.D. Mastrandrea, T.E. Bilir,

M. Chatterjee, K.L. Ebi, Y.O. Estrada, R.C. Genova, B. Girma, E.S. Kissel, A.N. Levy, S. MacCracken,

P.R. Mastrandrea, and L.L. White (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New

York, NY, USA, pp. 411-484.

412

Executive Summary ........................................................................................................................................................... 414

6.1. Introduction: Point of Departure, Observations, and Projections ........................................................................... 417

6.1.1. Changes in Physical and Chemical Variables .................................................................................................................................... 418

6.1.1.1.Temperature and Salinity ..................................................................................................................................................... 418

6.1.1.2.Carbon Dioxide-Induced Acidiﬁcation .................................................................................................................................. 418

6.1.1.3.Hypoxia ................................................................................................................................................................................ 418

6.1.1.4.Light and Nutrients .............................................................................................................................................................. 420

6.1.2. Historical and Paleo-Records ............................................................................................................................................................ 420

6.1.2.1.Historical Observations ........................................................................................................................................................ 420

6.1.2.2.Paleontological Records ....................................................................................................................................................... 421

6.2. Diversity of Ocean Ecosystems and Their Sensitivities to Climate Change ............................................................ 423

6.2.1. Pelagic Biomes and Ecosystems ........................................................................................................................................................ 424

6.2.2. Benthic Habitats and Ecosystems ..................................................................................................................................................... 424

6.3. Climate Change Impacts from Organism to Ecosystem .......................................................................................... 424

6.3.1. Temperature Effects .......................................................................................................................................................................... 427

6.3.1.1.Principles .............................................................................................................................................................................. 427

6.3.1.2.Microbes .............................................................................................................................................................................. 428

6.3.1.3.Macroalgae and Seagrasses ................................................................................................................................................. 429

6.3.1.4.Animals ................................................................................................................................................................................ 429

6.3.1.5.Ecosystems ........................................................................................................................................................................... 431

Box 6-1. An Atlantic Ocean Example: Long-Term Responses of Pelagic Organisms and Communities to Temperature .. 434

6.3.2. Carbon Dioxide Effects ..................................................................................................................................................................... 432

6.3.2.1.Principles .............................................................................................................................................................................. 436

6.3.2.2.Microbes .............................................................................................................................................................................. 439

6.3.2.3.Macroalgae and Seagrasses ................................................................................................................................................. 440

6.3.2.4.Animals ................................................................................................................................................................................ 440

6.3.2.5.Ecosystems ........................................................................................................................................................................... 441

6.3.3. Life in Hypoxia and Anoxia ............................................................................................................................................................... 443

6.3.3.1.Principles .............................................................................................................................................................................. 443

6.3.3.2.Microbes .............................................................................................................................................................................. 443

6.3.3.3.Animals and Plants .............................................................................................................................................................. 443

6.3.3.4.Ecosystems ........................................................................................................................................................................... 443

6.3.4. Mixed Layer Depth and Light Shaping Net Primary Production ........................................................................................................ 444

6.3.5. Concurrent Responses to Multiple Drivers ........................................................................................................................................ 445

6.3.5.1.Principles .............................................................................................................................................................................. 446

6.3.5.2.Microbes .............................................................................................................................................................................. 447

6.3.5.3.Animals and Plants .............................................................................................................................................................. 447

Table of Contents

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Ocean Systems Chapter 6

6.3.5.4.Ecosystems ........................................................................................................................................................................... 448

6.3.6. Food Web Consequences .................................................................................................................................................................. 448

6.3.7. Marine Reptiles, Mammals, and Birds ............................................................................................................................................... 448

6.3.7.1.Principles .............................................................................................................................................................................. 448

6.3.7.2.Field Observations ................................................................................................................................................................ 449

6.3.8. Summary and Conclusions ................................................................................................................................................................ 450

6.4. Human Activities in Marine Ecosystems: Adaptation Beneﬁts and Threats ............................................................ 451

6.4.1. Ecosystem Services ........................................................................................................................................................................... 452

6.4.1.1.Food from the Sea ................................................................................................................................................................ 452

6.4.1.2.Other Provisioning Services .................................................................................................................................................. 453

6.4.1.3.Climate Regulation and Extreme Events .............................................................................................................................. 453

6.4.1.4.Cultural Services ................................................................................................................................................................... 453

6.4.1.5.Supporting Services .............................................................................................................................................................. 453

6.4.2. Management-Related Adaptations and Risks ................................................................................................................................... 453

6.4.2.1.Ecosystem Management ...................................................................................................................................................... 453

6.4.2.2.Geoengineering Approaches ................................................................................................................................................ 454

6.4.2.3.Health Issues ........................................................................................................................................................................ 454

6.4.3. Conclusions ...................................................................................................................................................................................... 456

6.5. Projections of Future Climate Change Impacts through Modeling Approaches ..................................................... 456

6.5.1. Oceanic Primary Production .............................................................................................................................................................. 456

6.5.2. Higher Trophic Levels ........................................................................................................................................................................ 456

6.5.3. Ecosystems and Fisheries .................................................................................................................................................................. 457

6.5.4. Conclusions ...................................................................................................................................................................................... 459

6.6. Chapter Conclusions and Key Uncertainties ........................................................................................................... 461

6.6.1. Key Risks Related to Climate Change: Constraints on Ecosystem Services ....................................................................................... 461

6.6.1.1.Redistribution and Constraints on Microbial Functions and Primary Productivity ................................................................ 461

6.6.1.2.Warming-Induced Species Redistribution, Loss of Biodiversity, and Fisheries Catch Potential .............................................. 461

6.6.1.3.Expanding Hypoxia Affecting Marine Resources .................................................................................................................. 464

6.6.1.4.Constraints on Marine Calciﬁers and Associated Fisheries and Aquaculture due to Ocean Acidiﬁcation .............................. 464

6.6.1.5.Interactions of Climate-Related Drivers Exacerbating Impacts on Organisms, Ecosystems, and Their Services ..................... 465

6.6.2. Key Uncertainties .............................................................................................................................................................................. 465

References ......................................................................................................................................................................... 465

Frequently Asked Questions

6.1: Why are climate impacts on oceans and their ecosystems so important? ........................................................................................ 417

6.2: What is different about the effects of climate change on the oceans compared to the land,

and can we predict the consequences? ............................................................................................................................................ 426

6.3: Why are some marine organisms affected by ocean acidiﬁcation? ................................................................................................... 436

6.4: What changes in marine ecosystems are likely because of climate change? .................................................................................... 451

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Chapter 6 Ocean Systems

Executive Summary

Ocean ecosystems have responded and will continue to respond to climate changes of different rates, magnitudes, and durations

(virtually certain). Human societies depend on marine ecosystem services, which are sensitive to climate change (high confidence),

in particular the provisioning of food (fisheries and aquaculture) and other natural resources; nutrient recycling; regulation of global climate

including production of oxygen (O

) and removal of atmospheric carbon dioxide (CO

); protection from extreme weather and climate events;

and aesthetic, cultural, and supporting services. {6.3, 6.4, 6.5}

Climate change alters physical, chemical, and biological properties of the ocean (very high confidence). Oceanic drivers include

salinity, circulation, temperature, carbon dioxide (CO

), oxygen (O

), nutrients, and light. These drivers shape the physiological performance of

individual cells and organisms and ultimately determine ecosystem composition, spatial structure, and functioning. {6.1.1, 6.3}

The fossil record and present field and laboratory observations confirm links between key environmental drivers and responses

of ocean ecosystems to climate change (high confidence). For millions of years in Earth history, natural climate change at rates slower

than today’s anthropogenic change has led to significant ecosystem shifts (high confidence), including species emergences and extinctions

(high confidence). Contemporary multi-decadal natural climate variations associated with regional transient warming periods by 1°C have led

to fundamental restructuring of ecosystems and large socioeconomic implications (high confidence). {6.1.2, 6.3.1, 6.4}

Vulnerability of most organisms to warming is set by their physiology, which defines their limited temperature ranges and hence

their thermal sensitivity (high confidence). Temperature defines the geographic distribution of many species and their responses to climate

change. Shifting temperature means and extremes alter habitat (e.g., sea ice and coastal), and cause changes in abundance through local

extinctions and latitudinal expansions or shifts (very high confidence). Vulnerability is greatest in polar animals owing to their narrow temperature

ranges (medium confidence) and in tropical species living close to upper thermal limits (medium confidence). Although genetic adaptation occurs

(medium confidence), the capacity of present-day fauna and flora to compensate for or keep up with the rate of ongoing thermal change is

limited (low confidence). {6.3.1, 6.3.5, 6.5.2}

The warming-induced shifts in the abundance, geographic distribution, migration patterns, and timing of seasonal activities of

species (very high confidence) have been and will be paralleled by a reduction in their maximum body size (medium confidence).

This has resulted and will further result in changing interactions between species, including competition and predator-prey

dynamics (high confidence).

Numerous observations over the last decades in all ocean basins show global-scale changes including large-

scale distribution shifts of species (very high confidence) and altered ecosystem composition (high confidence) on multi-decadal time scales,

tracking climate trends. The distribution and abundance of many fishes and invertebrates have shifted poleward and/or to deeper, cooler waters

(high confidence). Poleward displacements of phyto- and zooplankton have occurred by hundreds of kilometers per decade (high confidence).

Some warm-water corals and their reefs have responded with species replacement, bleaching, and a decreased coral cover causing habitat loss

(high confidence). While marine reptiles such as turtles encounter direct effects of warming, impacts to seabirds and marine mammals are

mostly indirect through effects of warming on their prey (high confidence). {6.3.1, 6.3.7, 6.5, Boxes CC-CR, CC-MB}

In response to further warming by 1°C or more by the mid-21st century and beyond, ocean-wide changes in ecosystem properties

are projected to continue (high confidence).

Large irreversible shifts in the spatial distribution of species and seasonal timing of their

activities (feeding, growth, development, behaviors, and productivity) will have implications for species composition, and ecosystem goods and

services. {6.3.1, 6.4, 6.5, 6.6}

By the mid-21st century, the spatial shifts of marine species will cause species richness to increase at mid- and high latitudes

(high confidence) and to decrease at tropical latitudes (medium confidence), resulting in global redistribution of catch potential

for fishes and invertebrates, with implications for food security (medium confidence).

Animal displacements are projected to lead to

high-latitude invasions and high local extinction rates in the tropics and semi-enclosed seas. This will cause a 30 to 70% increase in the fisheries

yield of some high-latitude regions by 2055 (relative to 2005), a redistribution at mid-latitudes, but a drop of 40–60% in the tropics and the

Antarctic, based on 2°C warming above preindustrial values (medium confidence in the direction of trends in fisheries yields, low confidence in

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Ocean Systems Chapter 6

the magnitude of change). If a decrease in global net primary production (NPP) or a shift toward smaller primary producers occurs, the overall

fisheries catch potential may also decrease. {6.3.1-4, 6.4.1, 6.5.1-4}

Open ocean NPP is projected to fall globally depending on RCP scenario (medium confidence). The estimated decrease will occur

by up to 9% by 2100 under the RCP8.5 business-as-usual climate scenario (relative to 1990, low confidence). The oceans currently

provide about half of global NPP. Environmental controls on NPP include temperature, CO

, nutrient supply, and light (through cloud cover,

mixed layer depth), all of which will be altered (WGI AR5 Section 6.3). Present observations indicate increasing NPP at high (Arctic) latitudes

(medium confidence), projected to continue beyond 2100 (medium confidence). This increase is offset by a decrease at temperate and tropical

latitudes (medium confidence). Poor representation of shelf and coastal regions hamper projections in global NPP models for near-shore waters,

reducing confidence in global projections. {6.3.4, 6.5.1, Box CC-PP}

Large-scale processes and climatic feedbacks sustained by microbes (bacteria, archaea, unicellular algae, and protozoans) play

key roles in marine ecosystems (e.g., carbon and nitrogen (N

) fixation or nutrient recycling) and will be altered by climate

change (medium confidence). Identifying which microbial species, groups, and processes are being affected and how these will be altered is

difficult, as these organisms and their responses to environmental change are extremely diverse and often modulated by biological interactions or

changes in circulation and nutrient supply (limited evidence, low agreement). Warming will cause species-specific responses, such as enhancing

metabolic rates and exceeding thermal tolerances, which will affect abundance, distribution, and community structure. Warmer, CO

- and

nutrient-enriched coastal oceans may stimulate harmful algal blooms (medium confidence), and the redistribution of certain microbes causing

diseases such as cholera (medium confidence). {6.3, 6.4.2}

Rising atmospheric CO

over the last century and into the future not only causes ocean warming but also changes carbonate

chemistry in a process termed ocean acidification (WGI AR5 Sections 3.8.2, 6.4.4). Impacts of ocean acidification range from

changes in organismal physiology and behavior to population dynamics (medium to high confidence) and will affect marine

ecosystems for centuries if emissions continue (high confidence).

Laboratory and field experiments as well as field observations show a

wide range of sensitivities and responses within and across organism phyla (high confidence). Most plants and microalgae respond positively

to elevated CO

levels by increasing photosynthesis and growth (high confidence). Within other organism groups, vulnerability decreases with

increasing capacity to compensate for elevated internal CO

concentration and falling pH (low to medium confidence). Among vulnerable

groups sustaining fisheries, highly calcified corals, mollusks, and echinoderms are more sensitive than crustaceans (high confidence) and fishes

(low confidence). Trans-generational or evolutionary adaptation has been shown in some species, reducing impacts of projected scenarios (low

to medium confidence). Limits to adaptive capacity exist but remain largely unexplored. {6.3.2, Box CC-OA}

Few field observations conducted in the last decade demonstrate biotic responses attributable to anthropogenic ocean

acidification, as in many places these responses are not yet outside their natural variability and may be influenced by confounding

local or regional factors.

Shell thinning in planktonic foraminifera and in Southern Ocean pteropoda has been attributed fully or in part to

acidification trends (medium to high confidence). Coastward shifts in upwelling CO

-rich waters of the Northeast Pacific cause larval oyster

fatalities in aquacultures (high confidence) or shifts from mussels to fleshy algae and barnacles (medium confidence), providing an early

perspective on future effects of ocean acidification. This supports insight from volcanic CO

seeps as natural analogs that macrophytes

(seaweeds and seagrasses) will outcompete calcifying organisms. During the next decades ecosystems, including cold- and warm-water coral

communities, are at increasing risk of being negatively affected by ocean acidification, especially as ocean acidification will be combined with

rising temperature extremes (medium to high confidence, respectively). {6.1.2, 6.3.2, 6.3.5}

The expansion of hypoxic regions termed Oxygen Minimum Zones (OMZs) and anoxic “dead zones,” observed over the last 50

years and projected into the future under climate change, especially if combined with nutrient enrichment (eutrophication), will

constrain the habitat of O

-dependent organisms and benefit anaerobic microbes (medium confidence). Hypoxia tolerance varies

among species and is influenced by temperature, elevated CO

, food consumption, and O

demand (high confidence). Warming-induced

stratification limits the exchange of gases between water layers. Enhanced oxygen consumption by heterotrophic organisms depletes the oxygen

further, causing a community shift toward lower species richness and hypoxia-tolerant specialists. Under extreme hypoxia ecosystems are

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Chapter 6 Ocean Systems

dominated by microbes. These OMZs are also characterized by microbial removal of fixed nitrogen (denitrification), which can significantly

reduce the low-latitude nutrient inventories with implications for regional productivity. {6.3.3, 6.3.5}

The climate-change-induced intensification of ocean upwelling in some eastern boundary systems, as observed in the last decades,

may lead to regional cooling rather than warming of surface waters and cause enhanced productivity (medium confidence), but

also enhanced hypoxia, acidification, and associated biomass reduction in fish and invertebrate stocks.

Owing to contradictory

observations there is currently uncertainty about the future trends of major upwelling systems and how their drivers (enhanced productivity,

acidification, and hypoxia) will shape ecosystem characteristics (low confidence). {6.1.1, 6.3.2, 6.3.3, 6.3.5-6, Box CC-UP}

Environmental drivers acting simultaneously on ocean biota* often lead to interactive effects and complex responses (high

confidence). Interactions of temperature, ocean acidification, and hypoxia narrow thermal ranges and enhance sensitivity to temperature

extremes in organisms such as corals, coralline algae, mollusks, crustaceans, and fishes (high confidence). In primary producers, light and

individual nutrients can also interact with temperature and acidification. Combined warming and ocean acidification reduce calcification in

warm-water corals (high confidence). Ocean acidification will alter availability of trace metals (low confidence). (*The term biota encompasses

the organisms of a region, habitat, or geological period.) {6.3.2.2, 6.3.5, 6.5.2}

The combination and often amplification of global and regional climate change and local anthropogenic drivers result in

enhanced vulnerability of natural and human systems (high confidence).

Major regional and local drivers include fishing, pollution, and

eutrophication. {6.3.5, 6.4, 6.5}

The progressive redistribution of species and the reduction in marine biodiversity in sensitive regions and habitats puts the

sustained provision of fisheries productivity and other ecosystem services at risk, which will increase due to warming by 1°C or

more by 2100 compared to the present (high confidence).

Human societies respond with limited adaptive capacity. Socioeconomic

vulnerability is highest in developing tropical countries involving a risk of reduced supplies, income, and employment from marine fisheries

(high confidence). This emphasizes disparities in food security between developed and underdeveloped nations. {6.4.1, 6.5}

With continuing climate change, local adaptation measures (such as conservation) or a reduction in human activities (such as

fishing) may not sufficiently offset global-scale effects on marine ecosystems (high confidence). Effects of climate change will thus

complicate management regimes such as of marine protected areas once species undergo distributional shifts. This increases the vulnerabilities

of marine ecosystems and fisheries. {6.4.2.1}

Geoengineering approaches involving manipulation of the ocean to ameliorate climate change (such as nutrient fertilization,

binding of CO

by enhanced alkalinity, or direct CO

injection into the deep ocean) have very large environmental and associated

socioeconomic consequences (high confidence).

Some actually require purposeful alteration of ocean ecosystems for implementation.

Alternative methods focusing on solar radiation management (SRM) leave ocean acidification largely unabated as they cannot mitigate CO

emissions. {6.4.2}

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Ocean Systems Chapter 6

6.1. Introduction: Point of Departure,

Observations, and Projections

The oceans cover about 71% of Earth’s surface to an average depth of

700 m. Their importance for life on Earth, including humans, is vast (FAQ

6.1). Marine habitats display natural variability on various spatial and

temporal scales but a dearth of long-term observational data from the

vast open oceans limits our understanding of the causes and ecological

consequences of this variability. The available information indicates that

climate controls ocean temperatures, chemistry, circulation, upper ocean

stratification, nutrient supply, and sunlight exposure. These drivers affect

marine ecosystems through direct effects on organisms, amplified by

their changing interactions with other species. Food webs are modified by

changes in phytoplankton growth and the availability of live organisms or

their decomposing bodies, that is, debris or dissolved organic matter, as

food to (chemo-)heterotrophs (organisms gaining energy by feeding on

organic matter). Organismal responses lead to changes in biogeochemical

processes, such as the carbon cycle, and in biological diversity and the

services the oceans provide.

Some impacts of climate change on marine ecosystems and their services

were addressed in the IPCC Fourth Assessment Report (AR4): WGII

Chapters 4 to 6 (ecosystems, food, coastal areas), and regional chapters,

for example, 15 (polar regions) and 16 (small islands). The ecosystem

assessment in WGII AR4 Chapter 4 focused on terrestrial, coastal, and

marine systems, their properties, goods, and services. It emphasized the

difficulty in assessing future ecosystem responses as a result of ecosystem

complexity, different vulnerabilities of species, and ecosystem-specific,

critical thresholds associated with nonlinear responses to environmental

change. Focusing on terrestrial ecosystems, WGII AR4 Chapter 4 concluded

hat more than 2°C to 3°C warming above preindustrial levels causes

high extinction risks to 20 to 30% of present-day species (medium

confidence), paralleled by substantial changes in ecosystem structure

and functioning (high confidence). The authors projected that a wide

range of planktonic and benthic calcifiers will be impacted by ocean

warming (very high confidence) and acidification (medium confidence),

particularly in the Southern Ocean. They characterized sea ice and coral

reef biomes as highly vulnerable. Key uncertainties identified in AR4 were

the incomplete knowledge of ocean acidification (addressed in present

Section 6.3.2), synergistic effects and their mechanisms (Section 6.3.5),

biotic feedbacks to the climate system (Section 6.4), and the impacts

of interactions between climate change, human uses, and ecosystem

management (Section 6.4.2).

Much more than in previous IPCC reports (Figure 1-2), impacts on the

oceans are a focus in AR5. This allows for a more comprehensive

discussion of phenomena and impacts, as well as the associated

uncertainties and the levels of confidence in observed and projected

changes. The present chapter focuses on the general principles and

processes characterizing climate change impacts on ocean systems and

on the uses of these systems by human societies. For projections of

responses to climate change, the chapter also assesses our understanding

of underlying functional mechanisms causing change across all levels

of biological organization, from molecules to organisms to ecosystems.

As the ocean is a heterogeneous environment, the comparison of major

ocean regions is required to understand variability and differences in

key processes and carbon inventories (Box CC-PP, Figure 1). We discuss

the changes and variability in the ocean’s principal physical and chemical

properties and assess knowledge drawn from paleo- and historical to

present observations. We develop a conceptual framework for analyzing

Frequently Asked Questions

FAQ 6.1 | Why are climate impacts on oceans and their ecosystems so important?

Oceans create half the oxygen (O

) we use to breathe and burn fossil fuels. Oceans provide about 17% of the animal

protein consumed by the world’s human population, or almost 20% of that protein consumed by 3 billion people.

Oceans are home to species and ecosystems valued in tourism and for recreation. The rich biodiversity of the oceans

offers resources for innovative drugs or biomechanics. Ocean ecosystems such as coral reefs and mangroves protect

the coastlines from tsunamis and storms. About 90% of the goods the world uses are shipped across the oceans.

All these activities are affected by climate change.

Oceans play a major role in global climate dynamics. Oceans absorb 93% of the heat accumulating in the atmosphere,

and the resulting warming of oceans affects most ecosystems. About a quarter of all the carbon dioxide (CO

) emitted

from the burning of fossil fuels is absorbed by oceans. Plankton convert some of that CO

into organic matter, part

of which is exported into the deeper ocean. The remaining CO

causes progressive acidiﬁcation from chemical reactions

between CO

and seawater, acidiﬁcation being exacerbated by nutrient supply and with the spreading loss of O

content. These changes all pose risks for marine life and may affect the oceans’ ability to perform the wide range

of functions that are vitally important for environmental and human health.

The effects of climate change occur in an environment that also experiences natural variability in many of these

variables. Other human activities also inﬂuence ocean conditions, such as overﬁshing, pollution, and nutrient runoff

via rivers that causes eutrophication, a process that produces large areas of water with low oxygen levels (sometimes

called “dead zones”). The wide range of factors that affect ocean conditions and the complex ways these factors

interact make it difﬁcult to isolate the role any one factor plays in the context of climate change, or to identify

with precision the combined effects of these multiple drivers.

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Chapter 6 Ocean Systems

ffects on organisms and ecosystems and assess present knowledge

derived from experiments, field studies, and numerical model projections

mostly using Representative Concentration Pathways (RCPs) of climate

change scenarios to provide trajectories of climate change drivers (Moss

et al., 2010). Finally, we assess the implications of such changes for

ecosystem services, and identify plausible socioeconomic consequences.

Assessing climate change impacts on coastal systems is the topic of

Chapter 5. An integrative treatment of regional climate changes and

impacts in seven key ocean regions is the focus of regional Chapter 30.

Marine issues are also included in regional Chapters 22 to 29, with a

focus on polar oceans (Chapter 28) and small islands (Chapter 29). Topics

important to several chapters, such as ocean acidification, upwelling

systems, primary productivity, changes in biogeography, and coral reefs,

are discussed in joint assessments presented in the respective cross-

chapter boxes.

6.1.1. Changes in Physical and Chemical Variables

Trends in ocean conditions over the last 60 years reflect significant human

impacts beyond natural variability on temperature, salinity, dissolved

inorganic carbon and oxygen content, pH, and other properties of the

upper ocean (e.g., Pierce et al., 2012; Sen Gupta and McNeil, 2012; WGI

AR5 Section 3.8, Table 10.1). With climate change, marine ecosystems are

and will be exposed to rising temperature, ocean acidification, expansion

of hypoxic zones, and other environmental drivers changing concomitantly.

6.1.1.1. Temperature and Salinity

Over the last 39 years, oceans have warmed at average rates of >0.1°C

per decade in the upper 75 m and 0.015°C per decade at 700 m depth

(WGI AR5 Section 3.2.2, Figure 3.1). Trends differ regionally, seasonally,

and interannually (WGI AR5 Section 2.7; for ocean regions see Section

30.5 in the present volume). Temperature changes are particularly large

at El Niño-Southern Oscillation (ENSO) with high (3- to 4-year) and low

(5- to 7-year) frequencies, and on multi-decadal scales (>25 years,

Figure 6-1). The strongest warming trends are found at high latitudes

where most of the inter-decadal variability occurs, while tropical oceans

are dominated by interannual frequencies. Global climate models have

explored changes in different frequency domains, but their spatial

resolution is poor (WGI AR5 Sections 11.3.3, 12.4.7).

Temperature variations are often accompanied by changes in salinity.

Increased salinity results from reduced precipitation relative to evaporation,

for example, above the thermoclines (layer separating the upper mixed

layer from deeper water where temperature and density change rapidly

with depth) of subtropical gyres at mid- to low latitudes since 1950

(WGI AR5 Chapter 3). Decreased salinity due to enhanced precipitation

relative to evaporation has occurred at some tropical and higher latitudes,

exacerbated by sea ice melt (Durack et al., 2012). Both warming and

freshening cause enhanced density stratification, a trend projected to

continue into the 21st century (WGI AR5 Chapter 3, Section 11.3.3,

Figure 12.34; Helm et al., 2010). Mean sea surface temperature in

2090 will be 2.7°C warmer than in 1990 (RCP8.5; WGI AR5 Chapter 12;

Bopp et al., 2013).

6.1.1.2. Carbon Dioxide-induced Acidification

Rising carbon dioxide (CO

) concentrations in air (given as partial

pressures, pCO2, in µatm) cause increasing upper ocean CO

levels

(Watson et al., 2009). Starting from a preindustrial value of 280 µatm

atmospheric pCO

levels will have reached around 500 µatm by 2050

following the Special Report on Emissions Scenarios (SRES; IPCC, 2000)

and all RCPs (Moss et al., 2010; Meinshausen et al., 2011). By 2100 values

are projected to reach between 420 µatm and 940 µatm depending on

the RCP. The rise in pCO

causes ocean acidification (OA), measured as a

decline in water pH (negative log of proton concentration), accompanied

by a fall in both carbonate ion (CO

2–

) concentration and the saturation

states (Ω) of various calcium carbonates (CaCO

; Zeebe and Westbroek,

2003; WGI AR5 Section 3.8.2, Box 3.2, Chapter 6, Figure 6.29). Hence,

the seawater solubilities of three forms of CaCO

, namely calcite,

magnesium-calcite, and aragonite, increase. These minerals are important

components of shells and skeletons of many marine organisms (Section

6.3.2).

Ocean acidification occurs on a background of natural temporal and

spatial variability of pH, pCO

, and Ω. In the open ocean, the mean pH

(total scale, pH

) of surface waters presently ranges between 7.8 and 8.4

(WGI AR5 Section 3.8.2). In stratified mid-water layers, largely isolated

from gas exchange between surface waters and air, decomposition of

organic material leads to lowered oxygen (O

) and elevated CO

levels

(Paulmier et al., 2011) associated with lower pH values. The few existing

field data of sufficient duration, resolution, and accuracy (WGI AR5

Figure 3.18) show that trends in anthropogenic OA clearly deviate from

the envelope of natural variability (Friedrich et al., 2012). OA presently

ranges between –0.0013 and –0.0024 pH

units per year (WGI AR5

Section 3.8.2, Table 3.2, Box 3.2; Dore et al., 2009). Average surface

ocean pH has decreased by more than 0.1 units below the preindustrial

average of 8.17. By 2100 pH is expected to change by –0.13, –0.22,

–0.28, and –0.42 pH

units, at CO

levels of 421, 538, 670, and 936 ppm

under RCP2.6, 4.5, 6.0, and 8.5 climate scenarios, respectively (WGI AR5

Figure 6.28). The rate of acidification in surface waters varies regionally

and is 50% higher in the northern North Atlantic than in the subtropical

Atlantic (Olafsson, 2009). Salinity reduction caused by ice melt or

excess precipitation (Jacobs and Giulivi, 2010; Vélez-Belchí et al., 2010)

exacerbates OA by diluting the concentrations of substances acting as

buffers (Steinacher et al., 2009; Denman et al., 2011). At high sustained

concentrations the changes in ocean chemistry will take thousands

of years to be buffered by the natural dissolution of CaCO

from sediments

and tens to hundreds of thousands of years to be eliminated completely

by the weathering of rocks on land (Archer et al., 2009).

6.1.1.3. Hypoxia

The average dissolved oxygen concentration in the ocean is presently

162 µmol kg

–1

(Sarmiento and Gruber, 2006). Concentrations range from

over 500 µmol kg

–1

in productive Antarctic waters super-saturated with

oxygen (Carrillo et al., 2004) to zero in coastal sediments and in

permanently anoxic deep layers of isolated water bodies, such as the

Black Sea and the Cariaco Basin. Hypoxia results from oxygen depletion

in excess of supply as in stratified water bodies (Section 6.1.1.2). Vast

Oxygen Minimum Zones (OMZs) exist between less than 100 and more

419

Ocean Systems Chapter 6

0 10 20 30 °C

-2

2–3 years3–5 years

5–8 years

8–15 years

5–25 years25–40 years

0 10 20 30 °C

verage temperature between 1911 and 2011

(a) (b)

(c)

(d)

emperature range (historical maximum–minimum values)

0 10 20 30 40 50 (%)

Interannual variability of sea surface temperature

Global Average Temperature and

Sea Surface Temperature

Anomalies (GSST)

Southern Oscillation Index (SOI)

North Atlantic Oscillation (NAO)

Atlantic Multi-decadal Oscillation

(AMO)

Paciﬁc Decadal Oscillation (PDO)

Contribution to the time series variability (%)

20 30

40 50 60 70

80 90 100 0

Very high frequency (2–3 years)

High ENSO* frequency (3–5 years)

Low ENSO* frequency (5–8 years)

Decadal (8–15 years)

Bi-decadal (15–25 years)

Multi-decadal (25–40 years)

*ENSO = El Niño-Southern Oscillation

Figure 6-1 | Sea surface temperature variability between 1911 and 2011. (a) The sea surface temperature average for the period. (b) The temperature range calculated as the

difference between the maximum and minimum values for each grid component during the century. (c) The spatial distribution of variability by time scales (based on the

Extended Reynolds Sea Surface Temperature, NOAA, 2012) corresponds to the multi-decadal (25 to 40 years), bi-decadal (15 to 25 years), decadal (8 to 15 years), low ENSO (El

Niño-Southern Oscillation) frequency (5 to 8 years), high ENSO frequency (3 to 5 years), and very high frequency (2 to 3 years) scales. The summed variabilities from the same

2°x2° box in all six maps corresponds to 100% of the time series variability. (d) The spectral density of some of the most widely used climate indices, accumulated in the same

frequency windows. The total bar length (100%) corresponds to the cumulative variability of each time series between the 2 and 40 year frequency window. Climate indices were

obtained from the NOAA ESRL Physical Sciences Division website.

420

Chapter 6 Ocean Systems

han 900 m depths in Eastern Atlantic and Pacific tropical oceans. The

ecological literature applies the term hypoxia (see Section 6.3.3) to O

concentrations below 60 µmol kg

–

(estimated at about 5% of global

ocean volume; Deutsch et al., 2011). Pacific OMZs regularly reach oxygen

levels below 20 µmol kg

–1

(about 0.8% of global ocean volume; Paulmier

and Ruiz-Pino, 2009), lower than Atlantic ones. Suboxic waters at <4.5

μmol O

–1

occupy about 0.03% of the ocean volume, mainly in the

northeastern tropical Pacific (Karstensen et al., 2008).

OMZs are naturally present in many habitats including marine sediments,

but are also expanding due to anthropogenic influences. Over the past

50 years, open ocean O

concentrations have decreased by a mean rate

of 0.1 to >0.3 µmol kg

–1

(WGI AR5 Section 3.8.3; Stramma et al.,

2008). In some OMZs the rate has been much higher due to warming,

increased stratification, and rising biological O

demand (WGI AR5

Section 3.8.3). Long-term declines in O

by about 7 µmol kg

–1

per decade

have been documented at mid-water depths over much of the subarctic

North Pacific (Keeling et al., 2010). In coastal regions, extremely hypoxic

“dead zones” that exclude animal life, have increased from 42 reported

in the 1960s to more than 400 in 2008 and been attributed to high

oxygen demand from eutrophication, the local enrichment of nutrients,

resulting in organic matter loading and its decay as well as nitrous oxide

formation and release (Naqvi et al., 2000; Díaz and Rosenberg, 2008;

Zhang et al., 2010).

uture warming will likely accelerate the spread of hypoxic zones,

especially in temperate to sub-polar regions. Most models project

decreasing global ocean oxygen contents by 1 to 7% from present-day

concentrations in 2100 (Keeling et al., 2010; WGI AR5 Figure 6.30 under

RCP8.5), with a mean decline by 3.4% in 2090 compared to the 1990s

(Bopp et al., 2013). Warming and freshening of the surface layer will

increase stratification and reduce the depth of winter mixing. The

evolution of low O

zones will be linked to changes in fluvial runoffs

(e.g. Milly et al., 2008; see also Section 5.3.4.3), the wind regime (e.g.,

Vecchi and Soden, 2007), as well as the intensity, duration, and seasonal

timing of upwelling events (Snyder et al., 2003; see also Section 30.5.2).

The potential contributions of destabilized methane hydrates and bacterial

methane oxidation to exacerbate hypoxia and acidification at high

latitudes remain to be explored (Westbrook et al., 2009). Currently, there

is no consensus on the future volumes of hypoxic and suboxic waters

because of large uncertainties in potential biogeochemical effects and

in the evolution of tropical ocean dynamics due to both natural and

anthropogenic causes (WGI AR5 Section 6.4.5). While volumes with O

concentrations <80 µmol kg

–1

are projected to increase by several

percent, suboxic waters <5 μmol O

–1

may undergo a 30% increase

by 2100 compared to 2005 (low confidence; Bopp et al., 2013).

6.1.1.4. Light and Nutrients

Most models project that the mixed layer at the ocean surface (see

Figure 6-2) will become shallower in the coming decades through a

strengthening of the vertical density gradient (e.g., Sarmiento et al., 1998;

Sallée et al., 2013). Mean light levels encountered by phytoplankton are

set by incoming light from solar radiation, the depth of the mixed layer,

and the degree to which underwater light is attenuated by living and

non-living particles (Kirk, 1994). A shallower mixed layer will likely result

in the resident phytoplankton receiving higher mean underwater light

levels if the organisms are physically mixed through this stratum (Figure

6-2).

Enhanced, seasonally prolonged stratification (Holt et al., 2010), especially

in the tropics, the North Atlantic, the Northeast Pacific, and the Arctic

(Capotondi et al., 2012), will lead to decreased vertical transport of

nutrients to surface waters (Doney, 2010; Figure 6-2). River plumes

(Signorini et al., 1999), nutrient accumulation in the pycnocline as

reported for North Pacific waters (Whitney, 2011), human-induced

eutrophication, enhanced upwelling (Box CC-UP), and tidal mixing and

estuarine circulation in coastal oceans could partly compensate for the

projected reduction in nutrient supply in the oceans (limited evidence,

medium agreement).

6.1.2. Historical and Paleo-Records

6.1.2.1. Historical Observations

Ocean ecosystems are variable in time and space, and in a non-steady-

state, reflected in indices such as the North Atlantic Oscillation (NAO)

Index, the Atlantic Multi-decadal Oscillation (AMO), the Arctic Climate

Regime Index (ACRI), Pacific Decadal Oscillation (PDO), or the El Niño-

Southern Oscillation (ENSO) (WGI AR5 Box 2.5; Figure 6-1; Section 30.5).

Nutrient

supply

Nutrient

supply

Light

NOW FUTURE

Low oxygen mid-water

Carbonate solubility

Storms

Light

Dust

Warmer - fresher - acidiﬁed

Figure 6-2 | Projected alteration (magnitude and frequency) of oceanic ﬂuxes and

atmospheric events due to a changing climate in the coming decades. Ocean

properties will be altered from the sunlit surface layer to the mid-water stratum. In the

surface ocean, the depth of the mixed layer (solid horizontal line) will shallow

resulting in higher mean light levels. Increased density stratiﬁcation (i.e., a

strengthening sea water density gradient represented by the increasing thickness of

the solid horizontal line) will reduce the vertical supply of nutrients for

photosynthesizing organisms residing in the mixed layer. Anthropogenic CO

will

acidify, that is, lower the pH of the surface ocean (note this happens in a pH range

higher than 7 such that oceans will remain alkaline but less so due to acidiﬁcation).

The penetration of acidiﬁed waters to depth will result in a shallower depth (dashed

horizontal line) at which CaCO

structures, such as shells, dissolve. At depth, the

location of low-O

waters will progressively become shallower. In addition, changes in

storm activity and dust deposition will inﬂuence ocean physics and chemistry, with

consequent effects on ocean biota and hence ecosystems (courtesy of Reusch and

Boyd, 2013).

421

Ocean Systems Chapter 6

he combination of large, global data sets such as Reynolds, National

Center for Atmospheric Research (NCAR), International Comprehensive

Ocean-Atmosphere Data Set (ICOADS) with multi-decadal time series,

for example, near Hawaii (HOT), Bermuda (BATS), the Ligurian Sea

(DYFAMED), the Canaries (ESTOC), Kerguelen Island (KERFIX), Hokkaido

Island (KNOT), and Taiwan (SEATS) has provided data on the physical

and biogeochemical state of the oceans (Karl et al., 2003). These have

been augmented by the limited-term, high-resolution programs World

Ocean Circulation Experiment (WOCE) and Joint Global Ocean Flux Study

(JGOFS).

Historical data sets provide baseline information on ecosystem states

and document the responses of biota to both natural variability in the

ocean system and surface ocean warming since the 1970s (Figure 6-3;

Section 6.3.1). Such data sets are rare and regionally biased. Examples

include changes in geographic ranges of plankton and seasonal timing

(phenology) of different components of the ecosystem detected by the

Continuous Plankton Recorder (CPR: e.g., Edwards et al., 2001; Richardson

et al., 2006; Box 6-1) or multi-decadal shifts in pelagic ecosystems

(CalCOFI) including higher parts of the food chain such as sardines and

anchovies (Brinton and Townsend, 2003; Chavez et al., 2003; Lavaniegos

and Ohman, 2003; see also Section 6.3.1) and the skeletal archives of

long-lived organisms such as coralline algae (Halfar et al., 2011), bivalves

(Schöne et al., 2003), and corals (De’ath et al., 2009).

Systematic, long-term interdisciplinary observations using repeated,

highly calibrated measurements at a given field site are required to

capture high- and low-frequency events, for example, regime shifts

(abrupt changes between contrasting, persistent states of any complex

system; deYoung et al., 2008). Direct observations are complemented

by satellite remotely sensed data sets. Ocean color data (e.g., Coastal

Zone Color Scanner (1978–1986), Sea-Viewing Wide Field-of-View

Sensor (SeaWiFS, 1997–2010), and Moderate Resolution Imaging

Spectroradiometer (MODIS-AQUA, 2002 to the present); McClain, 2009)

provide estimates of chlorophyll concentrations (a proxy for phytoplankton

stocks and net primary production (NPP); Sections 6.2.1, 6.3.4; Saba et

al., 2011). Total chlorophyll cannot be measured from space; therefore,

the near-surface value (approximately one optical depth) is extrapolated

to whole water-column chlorophyll based on vertical distribution using

region-specific algorithms. Large uncertainties persist, as these estimates

reflect both phytoplankton stocks and their physiological status

(Dierssen, 2010; Behrenfeld, 2011). The approximately 15-year archived

time series of SeaWiFS is too short to reveal trends over time and their

causes. It is an example for the general issue that undersampling of

ocean phenomena in time and space limits our current ability to assess

present states, to distinguish effects of anthropogenic change from

natural variability, and to project future changes (Henson et al., 2010;

Beaulieu et al., 2013; Box CC-PP).

6.1.2.2. Paleontological Records

Paleontological records in marine sediments provide long-term, low-

resolution data on the spatial distributions of organisms and their

abundances from all ages and latitudes. This information can be readily

related to the concurrent shifts in multiple environmental properties

that are also recorded in these sediments. The records provide insights

nto shifts, expansions, and contractions of biogeographic ranges;

species extinctions and emergences; and changes in species abundance,

as well as the environmental forcings to which organisms respond.

Temporal trends reveal influences of temperature, hypoxia, CO

, and

food availability on organisms and ecosystems (Section 6.1.1; Figure

6-3).

Owing to insufficient resolution, the geological record often does not

allow the direct attribution of a biological change to a single driver or

the identification of various drivers and their relative importance. Support

for projections of future changes in present-day ecosystems and their

services is thus limited (low confidence; Sections 6.4, 6.5). Nonetheless,

information gained from the geological record is invaluable, as both

paleo and present climatic shifts share the same combination and sign

of environmental changes: increasing atmospheric CO

causing

warming and CO

enrichment in the surface ocean, leading to enhanced

stratification of the upper ocean and a decrease in dissolved O

(WGI

AR5 Chapter 3; Section 5.3). A combination of models (WGI AR5

Chapters 3, 6, 12) and geological data can be used to forecast future

impacts on ocean biota (medium confidence).

The last glacial-interglacial transition is associated with an average

increase in atmospheric CO

of approximately 1 µatm per century

between 18 and 10 thousand years before present (kyr BP) (WGI AR5

Chapter 5), a significantly slower increase than the approximately 90

µatm in the last century (WGI AR5 Chapters 5, 6). Consequently, the

average pH change of 0.002 pH units per century during the glacial-

interglacial transition is small relative to the ongoing anthropogenic

perturbation of >0.1 pH unit per century (WGI AR5 Section 3.8.2).

Overall the upper glacial ocean was more O

-rich than today’s ocean

(Jaccard and Galbraith, 2012) and between 0.7°C and 2.7°C colder, with

strong regional differences of up to 10°C cooling in the North Atlantic

and 2 to 6°C in the Southern Ocean (WGI AR5 Chapter 5, Table 5.2).

During warming from the glacial into the interglacial marine plankton

such as foraminifera, coccolithophores, diatoms, dinoflagellates, and

radiolarians showed marked poleward range expansion (high confidence;

see WGI AR5 Section 5.7; CLIMAP Project Members, 1976; MARGO

Project Members, 2009). Under the lower glacial CO

concentrations,

calcification in planktonic foraminifera was higher (limited evidence,

medium agreement).

The most prominent abrupt climate change periods in the recent

geological record, developing within 10 to 100 years, are associated

with Dansgaard-Oeschger (DO) and Heinrich events (WGI AR5 Section

5.7), which occurred repetitively during the last 120 kyr. Whereas the

atmospheric changes happened within a few decades, the sea surface

temperature in the North Atlantic changed by up to 5°C within decades

to centuries (WGI AR5 Section 5.7). Southern Ocean temperature changes

were slower (hundreds to thousands of years; Barker et al., 2009). The

cold phase of a DO event led to the migration of polar foraminiferal

species toward the equator, in the North Atlantic as far south as the

Iberian Peninsula (Martrat et al., 2004). Abrupt (approximately 100-

year) abundance changes in the Southern Ocean were associated with

latitudinal shifts in the Antarctic Circumpolar Current and associated

species (Barker et al., 2009) akin to modern changes in plankton range

due to warming (Box CC-MB, Box 6-1). During the DO warm phases the

Monsoon-driven Arabian Sea upwelling records show enhanced primary

422

Chapter 6 Ocean Systems

1000

500

1500

2000

planktic foraminifer

coccolithophore

benthic

foraminifer

benthic

foraminifer

coccolithophore

benthic foraminifer

coralline alga

–0.3

0.0

0.3

100

benthic foraminifer

planktic

foraminifer

Calciﬁcation

(g m

–2

–1

)

coral (Caribbean)

coral (Great Barrier Reef)

coral

Warm-water species (%)

1.5

1.6

1.7

1.8

–2

(μatm)

Normalized calciﬁcation

250

300

350

–0.5

LOSCAR model

GENIE model

0.0

0.5

Age (Million years)

55.155.355.555.7 1700 1800 1900 2000 2100

Calendar year

Environmental changes

Biotic responses

Extinctions

Migration

Calciﬁcation

Ocean

temperature

anomaly

tmospheric

Warm-water species (%) Number of species

Normalized calcite

volume

(°C)

Paleocene

–

Eocene Thermal Maximum

Industrialization

roxy reconstructions

irect measurement

55.155.355.555.7

1700 1800 1900 2000 2100

AMO index Normalized (°C)

currently no evidence for

climate-related extinction in the

marine record

Atlantic Multi-decadal Oscillation (AMO)

index

(mean detrended sea surface temperature

anomaly for the North Atlantic)

coralline alga

–1

0.3

–0.3

0.0

–0.3

0.3

AMO index

Figure 6-3 | Environmental changes (top) and associated biological responses (bottom) for the Paleocene–Eocene Thermal Maximum (PETM, left) and the industrial era (right).

The PETM represents the best geological analog for the future ocean because of its rapid environmental change. Episodes of largest environmental change are indicated with

darker bands. Note the different time scale between the two columns. Both time intervals are characterized by rapid warming both on land and in the ocean (modern: Wilson et

al., 2006 and PETM: Kennett and Stott, 1991) and increases in CO

(modern: Etheridge et al. 1996; Keeling et al., 2005 and PETM: Zeebe et al., 2009 (LOSCAR model); Ridgwell

and Schmidt, 2010 (Grid Enabled Integrated Earth System Model (GENIE model))). For the recent industrial era, the Atlantic Multi-decadal Oscillation (AMO; see Figure 6-1 and

Section 6.1.2.1) is shown to highlight an example of high-frequency sea surface temperature ﬂuctuations (Enﬁeld et al., 2001) and their inﬂuence on marine biota. Note the

species-speciﬁc calciﬁcation responses to climate change with decreases, increases, and high variability (coralline alga: Halfar et al., 2011; coral: Vásquez-Bedoya et al., 2012;

De’ath et al., 2013; PETM: Foster et al., 2013). While there was extinction during the PETM (Thomas, 2003), there is currently no evidence for climate-related extinction in the

marine record. Warming led to migration of warm-water species into previous cold-water habitats (modern: Field et al., 2006; PETM: Bralower, 2002). Pictures are examples of

organisms highlighting the processes in each panel, and are not to scale.

423

Ocean Systems Chapter 6

nd export production, reduced oxygenation, and denitrification, all

within approximately 200 years (Higginson et al., 2004).

The last time the atmospheric CO

content approached that of today

was during the Pliocene warm period (3.3 to 3.0 million years ago (Ma)),

with long periods of atmospheric CO

levels between 330 and 400 µatm

(Pagani et al., 2010; Seki et al., 2010) and equilibrated temperatures

approximately 2°C warmer than today (medium confidence; Haywood

et al., 2009; WGI AR5 Chapter 5). The Mid-Pliocene Warm Period saw a

poleward expansion of tropical planktonic foraminifera (high confidence;

Dowsett, 2007). Coccolithophores (Bown et al., 2004), corals (Jackson

and Johnson, 2000), and mollusks (Vermeij and Petuch, 1986) remained

unaffected with respect to rates of species extinction or emergences

compared to background rates.

Perhaps the best analog for the future ocean is the Paleocene-Eocene

Thermal Maximum (PETM, 55.3 Ma). The PETM was an event of warming

(Dunkley Jones et al., 2013), and ocean acidification (Zachos et al., 2005)

over millennia (Cui et al., 2011; Stassen et al., 2012) with increased

runoff and nutrients into the shelf ecosystems. Model simulations for

the PETM show 10 times lower rates of CO

input and hence ocean

acidification compared to today (medium confidence; Ridgwell and

Schmidt, 2010). Depending on the assumed rate and magnitude of the

release, models project pH declined by 0.25 to 0.45 units in PETM

surface waters and a reduction in surface ocean aragonite saturation

from Ω = 3 to Ω = 2 or even as low as 1.5 (Ridgwell and Schmidt, 2010).

Warming caused range expansions of warm-water taxa toward higher

latitudes (high confidence). The composition of plankton assemblages

changed both within and between phytoplankton groups (Gibbs et al.,

2006; Sluijs and Brinkhuis, 2009), possibly reflecting the warming trend

and/or changes in nutrient availability (Sections 6.2.2-3). There was no

bias in extinction toward more heavily calcifying species, possibly as slow

input led to minor surface water acidification. By contrast, benthic

foraminifera, the dominant deep water eukaryote, recorded up to 50%

extinction (Thomas, 2007). In contrast to sediment dwellers, more mobile

pelagic crustaceans (ostracods) did not show any significant change in

species composition (Webb et al., 2009). In shallow coastal waters,

calcareous algae and corals were replaced by symbiont-bearing benthic

foraminifera (medium confidence; Scheibner and Speijer, 2008).

The warm climates of the Mesozoic (251 to 65 Ma) led to a number of

anoxic events in the oceans (Jenkyns, 2010). In some cases, OMZs

expanded vertically, leading to anoxia in upper water layers (Pancost

et al., 2004). Some of the Cretaceous oceanic anoxic events were

associated with extinctions or increased species turnover (normalized

sum of originations and extinctions) of planktonic foraminifera and

radiolarians (30%). Such turnover was very small in other groups of

organisms (e.g., a maximum of 7% of coccolithophores; Leckie et al.,

2002). The attribution of these evolutionary changes to reduced O

tenuous as warming, changes in nutrient supply, and possibly ocean

acidification occurred concomitantly (Hönisch et al., 2012).

Global-scale collapse of marine ecosystems is rare, even in the geological

record. Some mass extinctions, in particular the Permian Period extinction

251 Ma, have been associated with large-scale inputs of carbon into

the atmosphere and ocean, with associated warming and deep-sea O

decline (Knoll et al., 2007; Kiessling and Simpson, 2011). The end-

ermian mass extinction preferentially affected reef organisms such as

corals and sponges resulting in a 4 Myr period without reef builders

(Kiessling and Simpson, 2011), and underscores that vulnerabilities

differ among organisms depending on anatomy, physiology, and ecology

(Knoll and Fischer, 2011). The rates of environmental change and any

potential acidification have not yet been accurately constrained for

these events.

Of the last 100 Myr, only the last 2 Myr had CO

levels of approximately

190 to 280 ppm, comparable to preindustrial values. Values like those

predicted for the mid and end of this century can solely be found in the

geological record older than 33 Ma, with large uncertainties in the

absolute numbers (WGI AR5 Section 5.3; Hönisch et al., 2012). That

marine biota thrived throughout high CO

times cannot imply that

marine organisms will remain unaffected in a future warm, high-CO

world. The key environmental issue of the 21st century is one of an

unprecedented rate of change, not simply magnitude, of CO

levels

(Hönisch et al., 2012). The current rate and magnitude of ocean

acidification are at least 10 times faster than any event within the last

65 Ma (high confidence; Ridgwell and Schmidt, 2010) or even 300 Ma

of Earth history (medium confidence; Hönisch et al., 2012). The slower

events in geological history provide robust evidence (high agreement)

for environmentally mediated changes in biogeographic ranges of fauna

and flora, their compositional changes, extinctions, and, to much lesser

degree, emergences (very high confidence). No past climate change event

perfectly parallels future projections of anthropogenic climate change,

which is unprecedented in evolutionary history. Existing similarities

indicate, however, that future challenges (Sections 6.1.1, 6.3.1-8) may

be outside the adaptive capacity of many organisms living in today’s

oceans (low to medium confidence).

6.2. Diversity of Ocean Ecosystems and Their

Sensitivities to Climate Change

Global-scale observation and modeling studies provide robust evidence

of present and future climate-mediated alterations of the ocean

environment (high agreement; Section 6.1.1; WGI AR5 Chapters 3, 6;

Bopp et al., 2013), which in turn impact ocean ecosystems (high

confidence; Boyd and Doney, 2002; Drinkwater et al., 2010; Hoegh-

Guldberg and Bruno, 2010). An assessment of present findings and

projections requires knowledge of the characteristics of ocean biota

and ecosystems and their climate sensitivity.

Life on Earth is diverse as a result of nearly 4 billion years of evolutionary

history. Marine microorganisms are the oldest forms of life and the most

functionally diverse; multicellular organisms are constrained to limited

functional abilities. Knowledge of overarching similarities across the

organism domains Archaea, Bacteria, and Eukarya (Woese et al., 1990)

or kingdoms Bacteria, Protozoa, Fungi, Plantae, Animalia, and Chromista

(Cavalier-Smith, 2004) would facilitate projections of climate impacts.

The phylogenetic and metabolic diversity of microbes (i.e., viruses,

archaea, bacteria, protists, and microalgae) sustains key ecosystem

processes such as primary production, CO

fixation and O

production,

the conversion of nitrogen into ammonia (N

fixation), and the use of

nitrate, sulfate, CO

, and metals (iron and manganese) in metabolism

instead of O

when it is absent. Microbes enhance the horizontal

424

Chapter 6 Ocean Systems

ransfer of genetic information between unrelated individuals, thereby

enhancing biodiversity (McDaniel et al., 2010). Microbes may respond

to climate change by exploiting their large diversity, undergoing species

replacements (Karl et al., 2001), and thereby sustain their biogeochemical

roles. Species replacements also occur among plants and animals, but

in most cases research has focused on their resilience, well-being,

abundance, survival, and conservation under climate change (FAQ

6.2).

6.2.1. Pelagic Biomes and Ecosystems

Pelagic organisms are key to biogeochemical processes in the ocean.

The base of the marine food web is the photosynthetic fixation of CO

by phytoplankton, a process termed (net) primary production (NPP;

Box CC-PP). Photosynthesis is controlled by light, temperature, inorganic

nutrients (CO

, nitrate, phosphate, silicate, and trace elements including

iron), and the density-dependent stability of the surface mixed-layer depth

(MLD) (Section 6.1.1; Figure 6-2; Sverdrup, 1953; González-Taboada and

Anadón, 2012). Environmental variability and the displacement of

organisms by ocean currents cause variability in phytoplankton

productivity, competitiveness, and natural selection (Margalef, 1978)

and result in changes in carbon sequestration (Box CC-PP; Figure 6-4).

Nutrient limitation leads to a decrease in NPP or chlorophyll levels and

a reduction in the amount of energy supplied to higher trophic levels,

including fish and invertebrates (high confidence; Ware and Thomson,

2005; Brander, 2007), affecting fishery yields (Cheung et al., 2008;

Friedland et al., 2012). The wide range of trophic structures in marine

food webs and the potentially nonlinear changes in energy transfer

under different NPP and temperature scenarios (Stock and Dunne, 2010)

hamper accurate projections of changes in higher trophic levels.

6.2.2. Benthic Habitats and Ecosystems

The ocean’s primary production is inextricably linked with benthic (sea

floor) communities via the biological pump (Figure 6-4), the chemical

exchange of nutrients and gases, and the existence of organisms with

both pelagic and benthic life history stages. Even in abyssal habitats, a

continuous rain of organic detritus serves as the primary source of carbon

nd energy. Therefore climate impacts on surface marine ecosystems

will impact even the deepest benthic communities, even if direct changes

to their physical habitat do not occur (Smith et al., 2009).

Benthic organisms living in shallow waters or the intertidal zone (where

they encounter temporary exposure to air) are exposed to widely

fluctuating and progressively changing means and extremes of

environmental variables, such as temperature, oxygen, CO

, salinity, and

sea level (WGI AR5 Chapters 3, 13; Sections 6.3.1-3, 6.3.5). Plants and

sessile or slow moving animals may be unable to escape from unfavorable

changes except by means of advection of fertilized eggs or planktonic

larvae. If climate change harms those species engineering benthic

habitats, the entire ecosystem may be impacted. This concerns those

ecosystem engineers, which form habitat from the structures they

produce (e.g., corals forming skeletons; Section 6.3.1) and those forming

habitat through their behavior (e.g., worms reworking and irrigating

sediment in a process termed bioturbation). Effects on both types of

ecosystem engineers (Sections 6.3.1-8) influence the regeneration of

nutrients and affect benthic-pelagic coupling.

6.3. Climate Change Impacts

from Organism to Ecosystem

Understanding climate-induced alterations in the functioning of

individual organisms, species populations, communities (assemblages

of various species), and ecosystems builds on studies in the laboratory,

in micro- and mesocosms (closed small- to medium-sized experimental

systems approximating natural conditions, holding selected biological

communities), and of biota or communities in the field as well as

modeling. These data inform us which taxonomic groups in what

regions are more susceptible to climate change (Boyd et al., 2011).

Empirical studies of marine organism and ecosystem sensitivities have

begun identifying the mechanisms and processes linking climate to

ecosystem changes (Drinkwater et al., 2010; Ottersen et al., 2010).

Changes in ecological community composition, species interactions, and

food web dynamics often build on organismal effects elicited by climate

forcing (e.g., Section 6.3.1.5; Boyd et al., 2010; Ottersen et al., 2010).

The underlying mechanisms respond to climate-related factors in a

hierarchy from organism (highest), tissue, cell to molecular (lowest)

Table 6-1 | To assess how a changing climate will alter the ocean’s biological pump (Figure 6-4) and determine the resulting biogeochemical feedbacks on global climate,

changes in a wide range of processes from cells to ocean basins, and from epipelagic to mesopelagic, must be quantiﬁ ed. This table illustrates the complexity of the integrated

knowledge platform needed to provide evidence of these biogeochemical ramiﬁ cations and thus the present limits to clear conclusions about climate-induced effects on the

biological pump (NPP = net primary production; C = carbon; TEP = transparent exopolymer particle; DOM = dissolved organic matter; POM = particulate organic matter).

Alteration of physiological

rates

Biogeographical changes /

community shifts

Altered foodweb structure:

trophodynamics

Changes to particle

dynamics

Biogeochemical changes /

climatic feedbacks

• NPP (Bopp et al., 2002, 2013)

• Particle solubilization through

bacterial ectoenzymes (Christian

and Karl, 1995)

• TEP production (Engel et al.,

2004)

• Microzooplankton grazing rates

(Rose et al., 2009)

• Microbial community structure

(Giovannoni and Vergin, 2012)

• Phytoplankton community

structure, e.g., biomes (Boyd and

Doney, 2002)

• Alteration of zooplankton biomes

(Beaugrand et al., 2009)

• Faunistic shifts at depth (Jackson

and Burd, 2001)

• Altered prey-predator linkages

(Lewandowska and Sommer,

2010)

• Faecal pellet geometry (Wilson et

al., 2008)

• C partitioning between DOM

vs. POM, e.g., TEP (Riebesell et

al., 2007)

• Sinking rates /seawater viscosity

(Lam and Bishop, 2008)

• Ballasting, e.g., calcite versus

opal (Klaas and Archer, 2002)

• Particle ﬂ ux /C sequestration

(Bopp et al., 2002)

• Shifts in elemental stoichiometry

of planktonic communities (Karl

et al., 2003)

• Remineralization rate; [O

hypoxia; nutrient resupply

(Gruber, 2011)

• Activity of the microbial loop;

vertical carbon export (Grossart

et al., 2006; Piontek et al., 2010)

425

Ocean Systems Chapter 6

Consumption

Aggregation

Fragmentation

Microbial

decomposition

Consumption

Sinking

Zooplankton

vertical

migration

Egestion

0–100 m

100–

500 m

>500 m

Physical mixing

of DOM

Ocean’s biological pump

Figure 6-4 | A schematic representation of the ocean’s biological pump, which will be inﬂuenced by climate change and is a conduit for carbon sequestration. It is difﬁcult to

project how the pump might be altered and whether it would represent a positive or negative feedback to climate change through the cumulative effects of affected processes,

surface to depth (Table 6-1): shifts in net primary production, ﬂoristic and faunistic community composition in the pelagic realm, and in grazing rates; alterations to the ballasting

of settling particles and the proportion of net primary production released as dissolved organic matter; modiﬁed bacterial enzymatic rates and particle solubilization; faunistic shifts

at depth. Note that the relative sizes of the organisms, particles, and particle building blocks are not presented to scale (modiﬁed from Buesseler et al. (2008) by J. Cook / WHOI).

Coccolithophorids

Radiolaria

Salps

Foraminifera

Diatoms

Other

Phytoplankton

Copepods

Euphausiids

Plankton Key

Salp pellets

Copepod pellets

Microzooplankton

mini-pellets

Aggregates (marine snow)

Aluminosilicates

Euphausiid pellets

Larvacean houses

Particle Key

Sinking

particles

426

Chapter 6 Ocean Systems

levels of biological organization (Pörtner, 2002a; Pörtner and Knust,

2007; Raven et al., 2012). Such knowledge aids the interpretation and

attribution to climate change of observed effects and is a major asset

for projections of future impacts.

The genetic and physiological underpinning of climate sensitivity of

organisms sets the boundaries for ecosystem response and provides

crucial information on sensitivities, resilience, and the direction and

scope of future change. As anthropogenic climate change accelerates,

a key issue is whether and how quickly organisms can compensate for

effects of individual or multiple drivers, by short-term acclimatization

or long-term evolutionary adaptation across generations. Evolutionary

adaptation depends on the genetic variation within a population,

from which the environment selects the fittest genotypes (Rando and

Verstrepen, 2007; Reusch and Wood, 2007). Genetic variation depends

on mutation rates, generation time, and population size (Bowler et al.,

2010). However, epigenetic mechanisms, such as modifications of the

genome by DNA methylation, can also influence fitness and adaptation

(Richards, 2006) and can be remarkably rapid as seen in terrestrial

ecosystems (Bossdorf et al., 2008). In plants and animals the rate of

evolutionary adaptation is constrained by long generation times, but

enhanced by high phenotypic variability and high mortality rates among

early life stages as a selection pool (e.g., Sunday et al., 2011). The limits

to acclimatization or adaptation capacity are presently unknown.

Frequently Asked Questions

FAQ 6.2 | What is different about the effects of climate change on the oceans

compared to the land, and can we predict the consequences?

The ocean environment is unique in many ways. It offers large-scale aquatic habitats, diverse bottom topography,

and a rich diversity of species and ecosystems in water in various climate zones that are found nowhere else.

One of the major differences in terms of the effect of climate change on the oceans compared to land is ocean

acidiﬁcation. Anthropogenic CO

enters the ocean and chemical reactions turn some of it to carbonic acid, which

cidiﬁes the water. This mirrors what is also happening inside organisms once they take up the additional CO

Marine species that are dependent on calcium carbonate (CaCO

), such as shellﬁsh, seastars, and corals, may ﬁnd it

difﬁcult to build their shells and skeletons under ocean acidiﬁcation. In general, animals living and breathing in

water like ﬁsh, squid, and mussels have between ﬁve and 20 times less CO

in their blood than terrestrial animals,

so CO

-enriched water will affect them in different and potentially more dramatic ways than species that breathe

in air.

Consider also the unique impacts of climate change on ocean dynamics. The ocean has layers of warmer and colder

water, saltier or less saline water, and hence less or more dense water. Warming of the ocean and the addition of

more freshwater at the surface through ice melt and higher precipitation increases the formation of more stable

layers stratiﬁed by density, which leads to less mixing of the deeper, denser, and colder nutrient-rich layers with

the less dense nutrient-limited layers near the surface. With less mixing, respiration by organisms in the mid-water

layers of stratiﬁed oceans will produce oxygen-poor waters, so-called oxygen minimum zones (OMZs). Large, more

active ﬁsh can’t live in these oxygen poor waters, while more simple specialized organisms with a lower need for

oxygen will remain, and even thrive in the absence of predation from larger species. Therefore, the community of

species living in hypoxic areas will shift.

State-of-the-art ecosystem models build on empirical observations of past climate changes and enable development

of estimates of how ocean life may react in the future. One such projection is a large shift in the distribution of

commercially important ﬁsh species to higher latitudes and reduced harvesting potential in their original areas.

But producing detailed projections, for example, what species and how far they will shift, is challenging because

of the number and complexity of interactive feedbacks that are involved. At the moment, the uncertainties in

modeling and complexities of the ocean system even prevent any quantiﬁcation of how much of the present

changes in the oceans are being caused by anthropogenic climate change or natural climate variability, and how

much by other human activities such as ﬁshing, pollution, etc.

It is known, however, that the resilience of marine ecosystems to adjust to climate change impacts is likely to be

reduced by both the range of factors and their rate of change. The current rate of environmental change is much

faster than most climate changes in the Earth’s history, so predictions from longer term geological records may not

be applicable if the changes occur within a few generations of a species. A species that had more time to adapt in

the past may simply not have time to adapt under future climate change.

427

Ocean Systems Chapter 6

owever, mass extinctions occurring during much slower rates of climate

change in Earth history (Section 6.1.2) suggest that evolutionary rates

in some organisms may not be fast enough to cope.

Comprehensive understanding of climate change effects on ecosystems

requires addressing the effects of individual drivers across organism taxa

(Sections 6.3.1-4), the integrated action of multiple drivers (Section

6.3.5), the consequences for food webs (Section 6.3.6), and the specific

effects on animals breathing in air (Section 6.3.7) and operating at the

highest trophic levels.

6.3.1. Temperature Effects

The effects of temperature on ecosystems largely result from organismal

responses. This requires that information on organisms’ thermal

sensitivities, limits, and functional properties is used to assess how

temperature changes have affected and will continue to affect species

distributions, abundances, diversity, trophic interactions, community

assemblages, risks of species extinctions, and ecosystem functioning.

rganisms also respond to temperature-driven changes in the physical

environment such as stratification, reduced sea ice cover, and freshening.

Ambient temperature interacts with other drivers such as ocean

acidification and hypoxia (Section 6.3.5). Ambient temperature plays a

more limited role for marine mammals and seabirds (Section 6.3.7).

.3.1.1. Principles

All organisms including marine ones have limited temperature ranges

within which they live and function. Organismal performance is related

to temperature by curves called thermal reaction norms (Figure 6-5),

which likely apply across all organisms (Chevin et al., 2010), from viruses

(Knies et al., 2006), bacteria (Ratkowsky et al., 1983), and phytoplankton

(Eppley, 1972; Thomas et al., 2012) to macroalgae and plants (Bolton

and Lüning, 1982; Müller et al., 2009; Vitasse et al., 2010) and animals

(Huey and Kingsolver, 1989; Angilletta, 2009). Heat tolerance thresholds

differ greatly between organisms and are hypothesized to be lowered

by rising organizational complexity and body size (Pörtner, 2002a,b).

Maximum heat limits of animals and plants are close to the maximum

Temperature range

denaturation

opt

anaerobiosis

loss of performance

and abundance

Scope for aerobic performance

Aerobic thermal

window

HighLow

Growing adults

Spawners

Eggs, early larvae

Juveniles

(a)

Thermal windows for animals: limits and acclimatization

(b)

Thermal window widths across life stages (ﬁshes)

Performance curve under normal conditions

Performance curve options under

elevated CO

or in hypoxic water or both

opt

Optimum temperature (performance maximum)

Pejus temperatures (limit to long-term tolerance)

Critical temperatures (transition to anaerobic metabolism)

Denaturation temperatures (the onset of cell damage)

acclimatization

and adaptation

Cold WarmCold Warm

threshold line

Figure 6-5 | Thermal specialization of an organism explains the why, how, when, and where of climate sensitivity. (a) The thermal tolerance range and performance levels of an

organism are described by its performance curve (exempliﬁed for an animal). Each performance (e.g., exercise, growth, reproduction) is maximal at its optimum temperature (T

opt

and becomes progressively constrained during cooling or warming. Surpassing the ﬁrst low- and high-temperature thresholds (T

; p, pejus: getting worse) means going into

time-limited tolerance. Once further cooling or warming surpasses the next low or high thresholds (T

; c, critical), oxygen availability becomes insufﬁcient and an anaerobic

metabolism begins. Denaturation temperatures (T

) are even more extreme and characterized by the onset of damage to cells and proteins. Horizontal arrows indicate that T

, T

and T

thresholds of an individual can shift, within limits, between summer and winter (seasonal acclimatization) or when the species adapts to a cooler or warmer climate over

generations (evolutionary adaptation). Under elevated CO

levels (ocean acidiﬁcation) and in hypoxic waters performance levels can decrease and thermal windows narrow

(dashed gray curves). (b) The width of the thermal range (horizontal arrows) also changes over time when an individual develops from egg to larva to adult and gains weight and

size. Blue to red color gradients illustrate the range between cold and warm temperatures (after Pörtner, 2002a, 2012; Pörtner and Farrell, 2008).

428

Chapter 6 Ocean Systems

temperature found in the warmest oceans (Figure 6-6). Knowledge of

reaction norms, thermal limits, and underlying mechanisms is most

advanced in animals (Pörtner et al., 2012; see also Section 6.3.1.4). Their

role in underpinning biogeography has not been explored systematically

in other organisms (e.g., Green et al., 2008), reducing the confidence level

in assessments of thermal impacts. In animals, changes in physiological

performances influence growth, body size, behavior, immune defense,

feeding, reproductive success, biogeography, phenology, and therefore

ecosystem structure and functioning. Shape and width of the curves

can shift through acclimatization and evolutionary adaptation (Figure

6-5a) and during life history (Figure 6-5b), with implications for the

distribution boundaries of species or populations (Section 6.3.1.5).

For any species, tracking the climate-induced displacement of tolerated

ambient temperatures by undergoing shifts in biogeographical ranges

to, e.g., higher latitudes during warming (Section 6.3.1.5; Figure 6-7) can

be understood as a simple mode of adaptation, implemented through

dispersal (e.g., of pelagic life stages), active movements (e.g., of migrating

adult fishes), or passive displacement (e.g., of early life stages or plankton

with drifting water masses). Conversely, fully completed acclimatization

or evolutionary adaptation (Figure 6-5) would involve shifting thermal

tolerance ranges and allow species to resist the temperature trend (e.g.,

warming) and to sustain fitness in their previous habitat.

6.3.1.2. Microbes

Temperature effects on growth, abundance, distribution, phenology, and

community structure of highly diverse microbes have large implications

for ecosystem functioning (Section 6.3; Box CC-PP). A warming ocean

may initially enhance the metabolic rates of microbes (Banse, 1991) and

stimulate their overall growth (Bissinger et al., 2008). Data from the

Continuous Plankton Recorder (Section 6.1.2) in the Northeast Atlantic

confirm that warming from 1960 to 1995 enhanced phytoplankton

growth (Edwards et al., 2001). Eventually, with warming, the thermal

tolerance of some groups will be challenged (Chevin et al., 2010), leading

to the replacement of species. This is reflected in increasing fractions of

smaller phytoplankton in warmer relative to colder waters (Morán et

al., 2010; Flombaum et al., 2013).

In response to transient warming, phytoplankton distribution in the

North Atlantic shifted poleward by hundreds of kilometers per decade

since the 1950s. Phenology of plankton in the North Atlantic was also

affected, with differences in sensitivity between groups (high

confidence; Section 6.3.1.5; Box 6-1). Coccolithophore blooms (Emiliania

huxleyi) in the Bering Sea were reported for the first time during the

period 1997–2000, probably in response to a 4°C warming, combined

with a shallower mixed layer depth, higher light levels and low

zooplankton grazing (Merico et al., 2004). Loss of multi-year Arctic sea

ice has had a profound effect on the diversity, structure, and function

of the epipelagic microbial assemblage (i.e., found in the layer into

which enough light penetrates for photosynthesis) (Comeau et al.,

2011), and further warming is likely to have even greater impacts

on the food web and on ecosystem services (medium confidence).

Warming may also have caused the southward range extension of

coccolithophores in the Southern Ocean in the 2000s (Cubillos et al.,

2007). However, further experimental and field observations (Giovannoni

and Vergin, 2012) are required to validate model projections (Taucher

Increasing complexity?

070605040 130120110100903020

Domain

Eukarya

Bacteria

Archaea

Multicellular

Unicellular

Cell number

Flagellata

Amoeba

Microalgae

Gram+ Bacteria

Flavobacteria

Cyanobacteria

Purple Bacteria

Ciliata

Thermotogales

Crenarchaeota

Euryarchaeota

Animals

Plants

Fungi

Group Tolerance range for growth

Temperature (°C)

ean SST = 17°C

Maximum SST = 41°C

aximal tempMinimal temp

Thermal window

ea surface

emperature (SST)

Figure 6-6 | Maximal values of temperature covered by various domains and groups of free-living marine organisms (bacteria to animals; domains and groups modiﬁed after

Woese et al., 1990). High organizational complexity is hypothesized to be associated with decreasing tolerance to heat and to enable an increase in body size which in turn,

decreases heat tolerance further (Sorokin and Kraus, 1962; Chevaldonné et al., 2000; Alker et al., 2001; Baumgartner et al., 2002; Pörtner, 2002a,b; Campbell et al., 2006; De

Jonckheere et al., 2009, 2011). In the domain Bacteria, the Thermotogales are less complex and most tolerant to high temperatures (Huber et al., 1986; Tenreiro et al., 1997;

Takai et al., 1999; Ventura et al., 2000; Abed et al., 2002). The highest temperature at which growth can occur is 122°C for hydrothermal vent archaea, seen under elevated

hydrostatic pressure in laboratory experiments (Kasheﬁ and Lovley, 2003; Takai et al., 2008).

429

Ocean Systems Chapter 6

nd Oschlies, 2011) of differential responses to warming by different

microorganisms.

6.3.1.3. Macroalgae and Seagrasses

Macrophytes in coastal waters (Chapter 5) cover 0.6% of the world’s

marine areas and supply about 2 to 5% of total oceanic production

(Smith, 1981; Charpy-Roubaud and Sournia, 1990; Field et al., 1998).

They have limited temperature ranges and are sensitive to temperature

extremes (high confidence), resulting in changes of photosynthesis,

growth, reproduction, and survival (following the principles of Figures

6-5, 6-6; and Harley et al., 2012), with consequences for their abundance,

distribution, and productivity. Ice retreat in polar areas leads to an

expansion of macroalgal distribution, for example, in the Antarctic

(Quartino et al., 2013).

Warm- versus cold-water-adapted species may have different sensitivities

to warming and show a range of responses in distribution shifts (Lima

et al., 2007). Temperate macroalgae with wide windows of thermal

tolerance acclimatize by shifting these windows following seasonal

temperature changes (Kübler and Davison, 1995). Antarctic and tropical

macroalgae are exposed to permanently low or high temperatures,

respectively, and have consequently specialized in a limited temperature

range, paralleled by a low acclimatization potential (Pakker et al., 1995;

Eggert et al., 2006; Gómez et al., 2011). Thus, Antarctic and tropical

macroalgae appear to be most vulnerable to warming (high confidence;

Short and Neckles, 1999). While observations in the tropics indicate that

seagrasses tolerate higher temperatures than seaweeds (Campbell et

al., 2006), an increase in maximum temperature by >1°C from 1988–

1999 to 2002–2006 (Section 30.5.3.1.5) led to increased seagrass shoot

mortality in the Mediterranean Sea (Marbà and Duarte, 2010). The

molecular basis of acclimatization and evolutionary adaptation, as well

as their limitation in relation to the climate regime, require further study

in the macrophytes.

6.3.1.4. Animals

The mechanisms shaping the thermal performance curve and, thereby,

an animal’s thermal niche have been explained by the concept of “oxygen

and capacity limited thermal tolerance” (OCLTT), applicable to marine

invertebrates and fishes (Pörtner et al., 2010; see also Figure 6-5a, FAQ

6.2). The temperature range at which animals can function best results

from optimal oxygen supply at minimal oxygen usage. At temperature

extremes, oxygen supply capacity becomes constrained in relation to

demand, and metabolism becomes thermally limited. Beyond upper and

lower temperature thresholds (T

, Figure 6-5a), growth, reproduction,

and other key functions decrease. These thresholds change during the

individual life cycle, and with body size. At large body size, limitations

in oxygen supply are exacerbated and heat tolerance limits shift to

lower temperatures.

Surpassing species-specific heat tolerance limits (Figure 6-5, T

) during

warming causes a reduction of abundance (Pörtner and Knust, 2007;

Katsikatsou et al., 2012), coral losses (Donner et al., 2005), shifts in the

seasonal timing of (zooplankton) biomass formation (Mackas et al.,

998; Schlüter et al., 2010), and changes in growth (Lloret and Rätz,

2000; Brunel and Dickey-Collas, 2010). During early life, owing to

incomplete development, or as adult spawners, owing to large body

size, animals may become more sensitive to warming because of

narrower thermal windows (Pörtner et al., 2008). This may cause high

vulnerability of winter-spawning Atlantic cod to warming winter to

spring temperatures (Table 6-2). In contrast, adult bigeye, bluefin,

and skipjack tuna spawn at high temperatures. They need to prevent

overheating by moving to cooler (deeper) waters (Lehodey et al., 2011).

Although temperature means are still most commonly used when

attributing responses of marine organisms to climate effects, temperature

extremes rather than means are most often mediators of effects (e.g.,

Easterling et al., 2000; Wethey et al., 2011; Wernberg et al., 2013; Figure

6-5). During heat exposure near the borders of the distribution range

(including the high intertidal or warming surface waters), reductions in

growth, activity, and abundance accompany even small (<0.5°C) shifts

in ambient temperature extremes (e.g., Takasuka and Aoki, 2006;

Pörtner and Knust, 2007; Nilsson et al., 2009; Neuheimer et al., 2011).

Local extinction events follow as a result of mortality or behavioral

avoidance of unfavorable thermal environments (Breau et al., 2011).

Shifted species distribution ranges follow temperature clines from high

to low, usually along latitudes, a lateral gradient at basin scale (Perry et

al., 2005; Poloczanska et al., 2013), or a vertical temperature gradient to

deeper waters (high confidence; Dulvy et al., 2008; Section 6.5.3; see

also Figure 6-5b, Box CC-MB).

Adopting OCLTT principles has enabled modeling studies to project

climate effects (Section 6.5), and paleo-studies to explain climate-induced

mass extinction events and evolutionary patterns in Earth history (Pörtner

et al., 2005; Knoll et al., 2007). For example, long-term observations show

that warming affects the body size of marine fishes (medium confidence).

Assessing effects of warming on body size may be complicated by

effects on the animal’s energy budget, the changing availability and

body size of prey species, community structure, species interactions, or

effects of fishing (Genner et al., 2010; Cheung et al., 2013a). Below the

thermal optimum, warming causes growth and weight-at-age of some

juvenile or younger fish populations to increase (e.g., Brunel and Dickey-

Collas, 2010; Neuheimer and Grønkjær, 2012). However, OCLTT predicts

that small individuals are more heat tolerant than large ones, in line

with observations of falling animal body sizes in warming oceans (Box

6-1; e.g., Daufresne et al., 2009). This trend is projected to continue into

the 21st century (medium to high confidence; Cheung et al., 2013a).

Thermal windows of fishes and invertebrates roughly match ambient

temperature variability (Figure 6-1) according to climate regime and

seasonality (Pörtner and Peck, 2010; Sunday et al., 2012). Sub-Arctic,

small, or highly mobile species are eurytherms. They function across a

wide temperature range, that is, they have wide thermal windows and

distribution ranges, at the expense of higher energetic costs and

associated lifestyles (Pörtner, 2002a, 2006). Conversely, high polar

species are stenotherms, that is, they have narrow thermal windows and

low energy demand lifestyles, making them sensitive to temperature

change. In a warming world, polar stenotherms will be marginalized,

with no possibility to escape to colder regions (high confidence).

However, extinction of polar species has not yet been reported. As

marine fishes and invertebrates in the Southern Hemisphere are

430

Chapter 6 Ocean Systems

adapted to less variable ocean temperatures than those in the Northern

Hemisphere (Jones et al., 1999; Figure 6-1), they may generally be more

vulnerable to warming extremes than Northern ones. Tropical species

(with thermal windows of intermediate width) live close to the highest

temperatures tolerated by marine animals (Figure 6-6). Vulnerability is,

therefore, highest for polar stenotherms, similar or lower for tropical,

and lowest for temperate species (high confidence).

Short-term shifts in thermal thresholds of an individual organism may

happen over days and weeks, such as during seasonal acclimatization.

Long-term shifts occur over many generations during evolutionary

adaptation of a population to cooler or warmer climates (Figure 6-5a;

Pörtner, 2006; Pörtner et al., 2008; Eliason et al., 2011). Both

acclimatization and adaptation involve adjustments in biochemical

characters (membranes, enzymes); however, the capacity to shift those

boundaries is limited and depends on the species and the prevailing

climate regime (Pörtner et al., 2008, 2012). Ocean acidification, hypoxia,

food availability, and stress affect those limits (Section 6.3.5; Figure

6-5a).

Local adaptation may reduce climate vulnerability at the species level,

by causing functional and genetic differentiation between populations,

thereby enabling the species to cover wider temperature ranges and

live in heterogeneous environments. Local adaptation on small spatial

scales is particularly strong in intertidal organisms (Kelly et al., 2012).

On larger scales, the widening biogeographic and roaming ranges of

Phenomenon Key drivers Mechanism / Sensitivity

Biogeography

orthward shift in the distribution of North Sea cod

(Gadus morhua) stocks between 1977 and 2001.

emperature Bottlenecks of high sensitivity during early life stages as well as

adult spawning stage in winter /early spring.

hift from sardines (Sardinops melanostictus) to

anchovies (Engraulis japonicus) in the western North

aciﬁ c observed between 1993 and 2003.

3,4

emperature Thermal windows of growth and reproductive output are

found at higher temperatures for anchovies than sardines, food

references of the competing species being similar.

Variable sensitivity of Paciﬁ c tuna species to the

vailability of dissolved O

Bigeye tuna routinely

reach depths where ambient O

content is below 1.5

l L

– 1

(≈ 60 µmoles kg

– 1

)

5, 6

Oxygen Oxygen transport via hemoglobin is adapted to be highly

fﬁ cient supporting high metabolic rates as needed during

feeding in the OMZ.

orthward movement of species and the conversion

of polar into more temperate and temperate

nto more subtropical system characteristics in

the European Large Marine Ecosystems between

958–2005.

, 8

arming and current advection Effects are attributed to climate change but may be inﬂ uenced

by nutrient enrichment and overﬁ shing.

Abundance

Increase in abundance of arctic boreal plankton

species, notably the copepods Calanus hyperboreus,

alanus glacialis and the dinoﬂ agellate Ceratium

arcticum between 1960 and 2000 in the

ewfoundland Shelf, Northwest Atlantic.

9,10

Temperature Temperature sensitivity of phyto- and zooplankter resulting from

cooling due to increased inﬂ ux of Arctic water.

A benthic ﬁ sh species, the eelpout (Zoarces

iviparus) at its southern distribution limit, the

German Wadden Sea, displayed abundance losses

during warming periods and rising summer extreme

emperatures between 1993 and 2005, with early

disappearance of the largest individuals.

Temperature Temperature extremes exceed organism’s thermal windows,

ith largest individuals being relatively less tolerant to high

temperature than smaller individuals.

ariable sensitivities to OA within and across animal

phyla (Figure 6-10b).

2 – 21

nthropogenic OA, sea water

acidiﬁ cation by elevated pCO

OMZs, upwelling areas, involving

anthropogenic ocean acidiﬁ cation.

owered extracellular (blood plasma) pH causing a lowering

of the rates of ion exchange and metabolism in muscle or liver

(hepatocytes) of vertebrates and invertebrates. High sensitivity

at reduced energy turnover in tissues and /or whole organism

by reduced ion exchange, use of more energy efﬁ cient transport

mechanisms, reduced protein synthesis, enhanced nitrogen

release from amino acid catabolism and protein degradation,

slower growth.

Phenology

Migration time of pink salmon (Oncorhynchus

gorbuscha) in Alaska is almost two weeks earlier in

2010s relative to 40 years ago.

Warming Rapid microevolution for earlier migration timing.

In the waters around the UK, during a period

of warming between 1976 and 2005, the

seasonal timing of biological events of all major

marine taxonomic groups (plant / phytoplankton,

invertebrate and vertebrates) advanced, on average,

by 0.31 to 0.43 days year

– 1

Warming Sensitivity to seasonal temperature changes as a result of

speciﬁ c thermal windows of different organisms.

Body size and growth

Asymptotic body sizes of different populations of

Atlantic cod (Gadus morhua) and Atlantic Herring

(Clupea harengus) are negatively related to

temperature.

24, 25

Warming At large body size, oxygen supply limitations are exacerbated

and the organism reaches its long-term heat tolerance limits at

lower temperatures, thus limiting the maximum body size that

can be reached.

Table 6-2 | Selected examples of species responses and underlying mechanisms to changing temperature, oxygen level and ocean acidiﬁ cation (OA). References are indicated by

superscript numbers and in the footnote.

1. Perry et al. (2005); 2. Pörtner et al. (2008); 3. Takasuka et al. (2007); 4. Takasuka et al. (2008); 5. Lehodey et al. (2011); 6. Seibel (2011); 7. Beaugrand et al. (2009); 8.

Philippart et al. (2011); 9. Johns et al. (2001);10. Greene and Pershing (2003); 11. Pörtner and Knust (2007); 12. Reipschläger and Pörtner (1996); 13. Pörtner et al. (2000); 14.

Vezzoli et al. (2004); 15. Langenbuch and Pörtner (2003); 16. Fernández-Reiriz et al. (2011); 17. Langenbuch and Pörtner (2002);18. Langenbuch et al. (2006); 19. Michaelidis et

al. (2005); 20. Pörtner et al. (1998); 21. Stumpp et al. (2012); 22. Kovach et al. (2012); 23. Thackeray et al. (2010); 24. Taylor (1958); 25. Brunel and Dickey-Collas (2010).

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Ocean Systems Chapter 6

orthern Hemisphere eurytherms into Arctic waters (Pörtner et al., 2008)

are supported by the differentiation into populations with diverse thermal

ranges, combined with high acclimatization capacity. By contrast, such

capacity is small in high polar, for example, Antarctic species (Peck et

al., 2010). Tropical reef fishes undergo rapid warm acclimation across

generations (Donelson et al., 2012) but some may approach animal heat

limits. The rates, mechanisms, and limits of thermal acclimatization and

evolutionary adaptation are poorly understood (low confidence).

6.3.1.4.1. Warm- and cold-water coral communities

Tropical corals live in shallow water and differ from most other animals

by hosting dinoflagellates (Symbiodinium sp.) in their tissues, which

provide the host with organic carbon from photosynthesis and with

nitrogen and enable the corals to build and sustain carbonate reefs (Box

CC-CR). High light, rapid salinity changes, and small increases in

temperature can trigger ”coral bleaching”, the loss of symbionts and tissue

color. In case of warming, early steps involve shifts in the photosynthetic

processing of light, generating Reactive O

Species (ROS) that may in turn

damage the symbionts (Hoegh-Guldberg and Smith, 1989; Glynn and

D’Croz, 1990; Jones et al., 1998; Hoegh-Guldberg, 1999). Mass bleaching

correlates with small temperature anomalies (+1°C to 2°C of the long-

term summer maximum, satellite observations), causing mortalities

(Goreau and Hayes, 1994; Strong et al., 2011) and decreasing coral

abundance, on average by 1 to 2% per year (high confidence; Bruno

and Selig, 2007; see also Box CC-CR; Section 30.5.6).

The degree of impact will depend on the coral reefs’ adaptability to

thermal stress and the interaction of multiple drivers (Meissner et al.,

2012; Teneva et al., 2012; see also Box CC-CR). Such capacity is

suggested by different heat tolerances among coral genera (Hoegh-

Guldberg and Salvat, 1995; Loya et al., 2001), the exchange of genetic

clades of Symbiodinium with more tolerant varieties (Baker, 2001; Jones

et al., 2008), as well as acclimatization phenomena (Howells et al., 2012).

Studies of the thermal sensitivity of deeper-living cold-water corals

(without endosymbionts) are scarce. One species, Lophelia pertusa,

responds to about 3°C warming with a threefold increase in metabolic

rate (Dodds et al., 2007), indicating a narrow thermal window in the

cold (cf. Pörtner, 2006).

6.3.1.5. Ecosystems

Heat exposure of ecosystem engineers may threaten the existence of a

whole ecosystem. During the last warm interglacial period equatorial

coral reefs deteriorated and retreated (Kiessling et al., 2012), a finding

emphasizing their thermal sensitivity (Veron et al., 2009) and showing

that warming oceans can reach temperatures well beyond the upper

heat limits of distinct animal groups and marine animals overall (Figure

6-6). In the present-day Great Barrier Reef, a large-scale survey found

diverse coral types along a climatic gradient, with no consistent

response to climatic drivers (Hughes et al., 2012). However, warm-induced

bleaching has contributed to the progressive decrease in live coral cover

observed over the last decades (De’ath et al., 2012; see also Box CC-CR;

Section 30.5.6).

ithin ecosystems, shifting competitive or trophic interactions, differential

risks for species extinctions and, thereby, scenarios of community-level

responses to temperature change (Urban et al., 2012; Milazzo et al., 2013)

can be traced back to changing differences in the performance of

participating animal species (Figure 6-7; e.g., Cairns et al., 2008; Harley,

2011; Pörtner, 2012). Knowledge is insufficient to assess interactions of

species from different domains, impeding a deeper understanding of

shifting distributions, abundances, community assemblages, and food

webs in space and time (low confidence in current understanding;

Parmesan and Matthews, 2005).

For example, in a coastal microcosm (small-scale, simplified experimental

ecosystem) resident heterotrophic bacteria were stimulated by warming

more than a laboratory-reared phytoplankter (Wohlers-Zöllner et al.,

2011). Also, high- to low-latitude transects in both the North and South

Atlantic revealed a shift between cold and warm waters, from photo-

autotrophs (gaining energy from photosynthesis) to chemo-heterotrophs

(Hoppe et al., 2002). Thermal stimulation of bacteria over phytoplankton

has biogeochemical implications, for example, microbially mediated

flow to the atmosphere might increase (Sarmento et al., 2010). The

principles and wider applicability of these findings require further

investigation (limited evidence, low agreement; Kirchman et al., 2009).

Observations of shifting distributions and phenologies, reproduction,

and range shifts of phytoplankton, zooplankton, other invertebrates,

fishes, and seabirds in pelagic and coastal marine ecosystems have at

least partly been attributed to temperature-mediated biological

responses (high confidence; see also Figure 6-8; Box 6-1; Box CC-MB).

In the North Atlantic as a key example, many biological events have

been occurring earlier in the year (robust evidence, high agreement;

Box 6-1; Section 30.5.1.1.1). Species richness has increased as a result

of shifts in ranges and abundances. In the Norwegian and Barents Seas,

a time series (1959–2006) of four commercial fish species and their

zooplankton prey showed that climate shapes population growth rates

through complex influences early in life, including direct temperature

effects on growth, indirect effects via the biomass of zooplankton prey,

and delayed feedback effects through predators (Stige et al., 2010).

Differential species responses to temperature and trophic amplification

were demonstrated to modify species interactions at five trophic levels:

primary producers (phytoplankton); primary, secondary, and tertiary

consumers (zooplankton, fishes, and jellyfishes); and benthic detritivores

(echinoderms and bivalves) (Kirby and Beaugrand, 2009). Also, the

responses of various plankton functional groups, such as diatoms,

dinoflagellates, and copepods, to warming are not synchronous, resulting

in predator-prey mismatches that carry over to higher trophic levels

(high confidence; Edwards and Richardson, 2004; Costello et al., 2006;

see also Figure 6-7a; Section 6.3.6). In the intertidal, warming-induced

changes in relative species ranges lead to shifts in dominance through

competitive interactions and to modifications in predator pressure

(Poloczanska et al., 2008; Harley, 2011). Trans-Arctic interchange of

species between Atlantic and Pacific has happened repeatedly in warm

periods of the Pleistocene (Dodson et al., 2007) and may occur again,

now facilitated by ballast transport by enhanced trans-Arctic shipping

(low to medium confidence).

Warming may increase the risk of disease outbreaks or parasite infections,

in marine organisms and ecosystems, and ultimately, humans (medium

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Chapter 6 Ocean Systems

confidence; Altizer et al., 2013; Burge et al., 2014). Some marine

pathogens and protist diseases are shifting their distribution poleward as

oceans warm (e.g., Baker-Austin et al., 2013; Burge et al., 2014). Climate

change may weaken the immune response of hosts, particularly fishes

and invertebrates, and increase their susceptibility to disease, as

observed during warming in coral reefs of the Pacific and Caribbean

(Harvell et al., 2009). Global outbreak frequencies of jellyfish aggregations

may follow rising sea surface temperatures (SSTs) (low confidence; Mills,

2001; Purcell and Decker, 2005), but evidence is inconclusive. Some

studies report an increasing trend (Brotz et al., 2012) and others do not

support this view (Condon et al., 2013).

In conclusion, organisms live in limited temperature ranges and are

sensitive to temperature extremes (very high confidence). Temperature

governs the biogeography, diversity, development, reproduction,

behavior, and phenology of marine species as well as the composition of

communities in both pelagic and benthic systems and the seasonal timing

of relevant processes (phenology) (very high confidence). Ecosystems

functioning at the coldest temperatures and warm adapted ones existing

at their upper thermal limits are more sensitive (medium confidence).

6.3.2. Carbon Dioxide Effects

Evidence for biological effects of ocean acidification stems from paleo-

observations (Section 6.1.2), few observations in the field (Section

6.3.2.5), studies at volcanic CO

seeps as natural analogs, and mostly

from short- to medium-term (hours to months) experiments in the

Latitudes (in °N)

Contraction

Expansion

Aerobic performance and productivity

Warm

adapted

Temperate

eurytherm

Temperate

Polar

Tropical

olar

tenotherm

oexistence ranges

Normal condition

, Hypoxia

Impact of photoperiod

High

North

Low

South

HighLow

(a)

Competition, predator/prey, phenologies of organisms in

different climate zones

(

patial dynamics during progressive warming

Temperature range

Under normal condition

Under elevated CO

and in

hypoxic water

Species abundance

Spatial dynamics during progressive warming

Phenology

shift

Seasonal temperature dynamics in low latitude

Temperature-dependent

time window

Temperature ranges of organisms in climate zones

easonal temperature dynamics in high latitude

Spring warming cue

Jan Dec

Warm

HighLow

Expansion Contraction

Cold

Performance

curve

Performance

range

Contraction

Expansion

Figure 6-7 | Role of thermal tolerance and performance of organisms at ecosystem level. (a) Thermal tolerance ranges (Figure 6-5) differ between species across polar, temperate,

and tropical climate zones, then overlap between coexisting species. Shifting temperatures and speciﬁc effects of additional drivers on the respective performance curves (dashed

lines) change the ﬁtness of coexisting species relative to each other as well as their temperature range of coexistence (after Pörtner and Farrell, 2008). Warming alters the timing

of seasonal activities (e.g., elicited by spring warming cues) to earlier, or can beneﬁt only one of two interacting species (e.g., in predator–prey dynamics or competition), causing

shifts in predominance. (b) During climate warming a largely unchanged thermal range of a species causes it to follow its normal temperatures as it moves or is displaced,

typically resulting in a poleward shift of the biogeographic range (exempliﬁed for the Northern Hemisphere; modiﬁed after Beaugrand, 2009). The polygon delineates the

distribution range in space and seasonal time; the level of gray denotes abundance. The Southern time window of tolerated temperatures shifts to earlier and contracts, while the

Northern one dilates (indicated by arrows). Species display maximum productivity in low latitude spring, wide seasonal coverage in the center, and a later productivity maximum

in the North. The impact of photoperiod (length of daily exposure to light) increases with latitude (gray arrow). Water column characteristics or photoperiod may overrule

temperature control in some organisms (e.g., diatoms), limiting northward displacement.

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Ocean Systems Chapter 6

0.00

0.04

0.08

Mean number of species per assemblage

2003–2005

2000–2002

1982–1999

50°N

60°N

1958–1981

50°N

60°N

(

a) Climate warming

(

b) Atlantic Multi-decadal Oscillation (AMO)

65°N

60°N

55°N

45°N

50°N

0°E 5°E 10°E 15°E5°W

Latitudes

72.15% of the geographical

cells have a temperature

regime between 9 and 10°C

Longitudes

(e) Warm-temperate pseudo-oceanic species

(f) Temperate pseudo-oceanic species

Annual mean

5-year

running mean

2003–2005

2000–2002

1982–1999

1958–1981

0.0

0.4

0.8

(d) Temperature regime (1988–2005)

11.5

10.5

9.5

8.5

0°E 5°E 10°E 15°E5°W

5°N

60°N

55°N

45°N

50°N

Latitudes

20.25% of the geographical

cells have a temperature

regime between 9 and 10°C

Longitudes

11.5

10.5

9.5

8.5

−

−15

−

−20

Index of temperature change

(AMO; 17.5%)

Index of temperature change

(climate change; 30.5%)

SST (in °C)

−

−15

−

−20

Figure 6-8 | Multi-decadal changes in ecosystem structure in the Northeast Atlantic driven by warming from both anthropogenic climate change and natural climate variability.

(a) Index of temperature change over the North Atlantic (31°N to 65°N and 99°W to 11°E) reﬂecting climate change. This index is the ﬁrst principal component (i.e., explaining

30.5% of observed variability) based on a principal component analysis (PCA) performed on sea surface temperature. (b) Index of temperature change (17.5% of observed

variability) reﬂecting the Atlantic Multi-decadal Oscillation (AMO). The index is the second principal component. (c, d) Observed mean annual sea surface temperature in the

North Sea during 1960–1981 (c) and 1988–2005 (d). The location of the critical thermal boundary (9°C to 10°C) is indicated by “+.” (e) Long-term changes in the mean number

of warm-temperate pseudo-oceanic species from 1958 to 2005. (f) Long-term changes in the mean number of temperate pseudo-oceanic species from 1958 to 2005. The period

1958–1981 was a period of relative stability and the period 1982–1999 was a period of rapid northward shifts, indicating that the abrupt ecosystem shift observed in the North

Sea was part of a large-scale response of the zooplankton biodiversity to warming temperatures (see a–d). Average values are below 1 because they are annual averages. Note

that the color bar is 10-fold smaller for warm-temperate pseudo-oceanic species because these species are less frequently observed than their temperate counterparts. Panels (a)

and (b) from Edwards et al. (2013), and (c)–(f) from Beaugrand et al. (2008, 2009).

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Chapter 6 Ocean Systems

Box 6-1 | An Atlantic Ocean Example:

Long-Term Responses of Pelagic Organisms and Communities to Temperature

Long-term observations (Sections 6.1.2, 30.5.1.1.1) encompassing the pelagic Northeast Atlantic over a 50-year period and longer

(Figures 6-8, 6-9) show changes in the seasonal abundance of phytoplankton, rapid northerly displacements of temperate and

subtropical zooplankton (e.g., calanoid copepods) and phytoplankton (e.g., dinoﬂagellates and diatoms), and the resulting changes in

the ecosystem functioning and productivity (high conﬁdence; Edwards et al., 2001; Beaugrand et al., 2002; Edwards and Richardson,

2004). The range limit of warm water copepods shifted by 10° north since 1960 (Beaugrand et al., 2009), with attendant mismatch in

the seasonal timing of trophic levels (predators and prey) and functional groups (Edwards and Richardson, 2004). Modes of climate

variability reﬂected in climate indices like the Northern Hemisphere Temperature (NHT) and the North Atlantic Oscillation (NAO) over

multi-decadal periods accompanied these changes (Figure 6-1). In cooler regions, increased phytoplankton activity caused by warming

favored growth, resulting in the observed increase in phytoplankton biomass, whereas a decrease in nutrient supply would have

prevented growth in warmer regions and caused a decrease in biomass (Richardson and Schoeman, 2004; see also Section 6.3.4).

Hinder et al. (2012) attributed a recent decline in North Sea dinoﬂagellates relative to diatoms to warming, increased summer

windiness, and thus water column turbulence. The ecosystem response to natural climate variability in the past provides a glimpse

into the climate-induced changes of the near future (Figure 6-9).

Temperature change (

Increasing magnitude

of change

1910 197019501930

2000 2020 204020302010

1.0

0.8

0.6

0.4

0.2

0.0

(a) Observed temperature variability

in North Atlantic

(b) Projected temperature change

(RCP8.5) in North Atlantic

Observed temperature

Projected temperature

Values beyond historical observations

Trend curves / Schematic for changes

Temperature maxima reached in the 1940s

parameters (see table on the right)

(d) Sketch of projected changes in

parameters (see table on the right)

Parameter sketched

by curves (c) and (d)

How parameter changes

Biogeography Poleward expansion and shift

Phenology Timing variability

Community composition Species turnover

NPP (mg C m

–2

day

–1

) Net primary productivity

Zooplankton (t km

–2

) Biomass and latitudinal shift

Fish stocks (t km

–2

)Biomass and latitudinal shift

Figure 6-9 | Schematic depiction of observed effects of approximately 1°C ocean warming in the northern North Atlantic driven by climate variability (a,c) versus

effects expected from anthropogenic climate change (b,d). (a) Transient warming and cooling associated with Atlantic Multi-decadal Oscillation (AMO) variability

(Drinkwater, 2006), based on the Kola Section temperatures (0 to 200 m; Stations 3 to 7, 71.5° to 72.5°N, 33.5°E) in the Barents Sea obtained from

http://www.pinro.ru and ﬁltered using a 20-year running mean. Similar trends occurred across most of the northern North Atlantic although the amplitude and timing

of the peaks and troughs varied spatially. (b) Warming driven by climate change for the same region (Representative Concentration Pathway 8.5 (RCP8.5) simulations

averaged from Coupled Model Intercomparison Project Phase 5 (CMIP5) models, computed as the mean over the upper 200 m in the grid box (2.5° × 2.5°) centered at

71.25°N and 33.75°E). (c) Warming and subsequent cooling in the northern North Atlantic during the period shown in (a) resulted in complex multi-faceted changes

(shown schematically) in net primary production (NPP), zooplankton biomass, and ﬁsh stock abundances. There was a general poleward shift and range expansion of

many commercial (e.g., Atlantic herring, Atlantic cod, haddock) and non-commercial species, reversed during the subsequent cooling period. Poleward shifts in

spawning areas (e.g., Atlantic cod) were also reversed as the waters cooled. Shifts in seasonal timing (phenology) and community composition were inﬂuenced by

earlier arrivals and later retreat of migratory ﬁsh (not shown). For more details see Drinkwater (2006). (d) Projected effects of climate mediated warming on northern

sub-polar and polar biota based on model projections of altered NPP (Bopp et al., 2013), and of the range shift of exploited ﬁshes and invertebrates (Cheung et al.,

2009, 2013a). The projected trends in (d) will differ with latitude, for example, decreased NPP at lower latitudes and no signiﬁcant change to NPP in temperate waters

(Bopp et al., 2013). Higher NPP supported and is projected to support higher trophic levels at high latitudes (c,d; Section 6.3.4). Note that climate variability will be

superimposed on anthropogenic warming (b; see Figures 6-1, 6-8a,b). Dashed lines indicate projected changes to continue beyond the range of historical observations.

Continued next page

435

Ocean Systems Chapter 6

Box 6-1 (continued)

In regions of high vulnerability to climate, mild warming can trigger rapid and substantial ecosystem shifts, offering a way to anticipate

future changes (Figure 6-9). In line with the increased understanding of physiology (Section 6.3.1.1), warming in the temperate to

polar North Atlantic was paralleled by a reduction in the average body lengths of about 100 copepod species, from 3 to 4 mm to 2 to

3 mm (Beaugrand et al., 2010). Warming also correlated with an increase in species richness among copepods and within the dinoﬂa-

gellate genus Ceratium. In diatoms, which are major contributors to carbon export (Armbrust, 2009), warming and decreasing annual

variability in SST resulted in lower diversity, smaller size, and reduced abundance (Beaugrand et al., 2010). Morán et al. (2010) found

that temperature alone explained 73% of the variance in the contribution of small cells (picophytoplankton) to total phytoplankton

biomass in the eastern and western temperate North Atlantic from –0.6 to 22°C. More recently, Marañón et al. (2012) analyzed data

from polar, sub-polar, and tropical regions and suggested that nutrient availability may inﬂuence cell size more than temperature.

The ecosystem regime shift observed in North Sea plankton in the late 1980s involved an increase in phytoplankton stocks and

changes in species composition and abundance among holozooplankton (animals that are planktonic for their entire lifecycle) (Reid

et al., 2001; Kirby and Beaugrand, 2009; Kirby et al., 2009; Lindley et al., 2010). This shift was paralleled by the northward propagation

of a critical thermal boundary (CTB, i.e., the boundary of the sub-polar gyre) between the temperate and the polar biomes (Beaugrand

et al., 2008; see also Box CC-PP, Figure 1). Warming to above the CTB coincided with pronounced and large-scale variations in

phytoplankton productivity, an increase in calanoid copepod diversity (Beaugrand et al., 2008) and herring abundance (Schlüter et al.,

2008), a reduction in the mean size of calanoids, and a decrease in the abundance of southern Atlantic cod populations in the North

Atlantic Ocean (e.g., the North Sea; Pörtner et al., 2008; Beaugrand et al., 2010). These patterns also extend to the southern North

Sea, where elevated salinities and average warming by 1.6°C, both in summer and winter between 1962 and 2007, expanded the

time window for growth of microalgae and possibly supported the invasion and increase in numbers of warm-adapted siliciﬁed

diatoms (Wiltshire et al., 2010). Recent ﬁndings indicate a regime shift in the Bay of Biscay and the Celtic and the North Seas in the

mid to end 1990s (Luczak et al., 2011). Changing plankton composition and changing abundances of both sardine and anchovies

(Raab et al., 2013) paralleled stepwise warming.

Northward range extensions or redistributions in ﬁshes were largest along the European Continental shelf and attributed to regional

warming, for example, by 1.0°C from 1977 to 2001 in the North Sea, with winter warming being closely correlated with the shift of

Atlantic cod (Perry et al., 2005; see also Section 6.3.1). Similar trends were observed due to warming by 1°C to 2°C in the waters

south and west of Iceland during the past 15 years (Valdimarsson et al., 2012). In the Northwest Atlantic Arctic and sub-Arctic, winter

and spring warming caused expansion of the area matching the thermal optimum of Atlantic salmon at 4°C to 8°C and caused

greater growth (Friedland and Todd, 2012). Pelagic sardines and anchovies entered the North Sea in the early to mid-1990s, after

about 40 years of absence, in response to intensiﬁed NAO and AMO (Alheit et al., 2012). Red mullet and bass extended into western

Norway; Mediterranean and northwest African species extended to the south coast of Portugal (Brander et al., 2003; Beare et al.,

2004; Genner et al., 2004; see also Section 30.5.1.1.4).

In the Northwest Atlantic cooling and freshening occurred during the late 1980s to early 1990s and seemed to have the opposite

effect, as capelin and their predator, Atlantic cod, shifted farther south (Rose and O'Driscoll, 2002). Between the early 1990s and mid-

2000s in the Northwest Atlantic sub-polar gyre, phytoplankton biomass increased, due to warming. At the same time, Arctic copepod

species became more abundant, due to increased inﬂux of Arctic water (Head and Pepin, 2010). Although temperatures have risen on

the Newfoundland Shelf (Colbourne et al., 2011), capelin and cod remain scarce for reasons probably unrelated to climate (DFO,

2011a,b). Farther south, Arctic freshwater inﬂows caused freshening and increased stratiﬁcation of the area around the Gulf of Maine

throughout the 1990s, resulting in enhanced phytoplankton abundance, a larger and later fall bloom, increased abundance of small

copepods, and a decrease in the large copepod Calanus ﬁnmarchicus (deYoung et al., 2004; Pershing et al., 2005, 2010). Various ﬁsh

species showed poleward shifts in distribution (Table 6-2) that were associated with reduced survival of larval cod (Mountain and

Kane, 2010) and fewer right whale calves (Greene et al., 2003), but increased herring abundance (Greene and Pershing, 2007).

436

Chapter 6 Ocean Systems

laboratory or field, exposing organisms to projected future CO

levels

(Sections 6.3.2.1-4). A surging number of studies is providing evidence

that rising CO

levels will increasingly affect marine biota and interfere

with ecological and biogeochemical processes in the oceans (high

confidence; FAQs 6.2, 6.3).

6.3.2.1. Principles

The absorption of rising atmospheric CO

by oceans and organisms

changes carbonate system variables in the water and in organism internal

fluids, that is, the relative proportions of CO

, carbonate, bicarbonate,

and hydrogen ions (pH). Internal pH must be tightly controlled, as some

processes, such as calcification, release protons thereby affecting pH and

as other biochemical processes are pH sensitive. Accumulation of CO

and the resulting acidification can also affect a wide range of organismal

functions, such as membrane transport, calcification, photosynthesis in

plants, neuronal processes in animals, growth, reproductive success, and

survival. Effects translate from organism to ecosystem.

The capacity of organisms to resist and compensate for the CO

-induced

acidification of internal fluids depends on acid-base regulation, that is,

the capacity of ion exchange to accumulate bicarbonate internally, an

aspect unexplored in many phyla (low to medium confidence; Figure

6-10a; e.g., animals: Heisler, 1986; Claiborne et al., 2002; Pörtner, 2008;

phytoplankton: Taylor et al., 2011; see also FAQ 6.3).

In unicellular microbes the regulation of intracellular pH may play a key

role in modulating CO

responses (Taylor et al., 2011). Findings in

invertebrates and fish indicate an additional role for extracellular pH

(Figure 6-10a); effective pH values may vary between species. Organisms

pre-adapted to elevated CO

may minimize the decrease in pH (acidosis).

They may also modify their sensitivity such that they respond less or

not at all to the acidosis. Recent evidence, however, emphasizes a role

for acid-base regulation in a natural low-pH setting. Between two

urchin species, only the one successful in maintaining its setpoints of

extracellular pH is able to settle close to volcanic CO

seeps (Calosi et al.,

2013). Compensating for the acidosis may cause increased energy demand

and respiration rates. In general, such capacity rises with metabolic

energy turnover, for example, it is higher in more active marine animals,

such as fishes, cephalopods, and pelagic copepods, and in mobile coastal

crabs compared to sessile species (Pörtner et al., 2005, 2011; Ishimatsu

et al., 2008; Melzner et al., 2009; Ishimatsu and Dissanayake, 2010; see

also Table 6-3). This matches the sensitivity distribution seen among

animals at the phylum level (medium confidence; Figure 6-9b).

Some species have lower metabolic rates in response to acidosis (Pörtner

et al., 1998; Michaelidis et al., 2005; Pörtner, 2008; Liu and He, 2012;

Navarro et al., 2013); others display increased energy turnover and food

ingestion rates, possibly indicating a capacity to resist acidification effects

(Parker et al., 2011; Saba et al., 2012). The effects of the acidosis on

various processes relevant to fitness may explain changes in whole-

organism energy demand, probably paralleled by modified ion exchange,

protein synthesis, and growth and feeding rates. The magnitude of effect

depends on the CO

concentrations reached (Figure 6-10b).

The internal formation of carbonate from bicarbonate is essential to

calcification, which is the formation of solid CaCO

in internal or external

Frequently Asked Questions

FAQ 6.3 | Why are some marine organisms affected by ocean acidiﬁcation?

Many marine species, from microscopic plankton to shellﬁsh and coral reef builders, are referred to as calciﬁers,

species that use solid calcium carbonate (CaCO

) to construct their skeletons or shells. Seawater contains ample

calcium but, to use it and turn it into CaCO

, species have to bring it to speciﬁc sites in their bodies and raise the

alkalinity (lower the acidity) at these sites to values higher than in other parts of the body or in ambient seawater.

That takes energy. If high CO

levels from outside penetrate the organism and alter internal acidity levels, keeping

the alkalinity high takes even more energy. The more energy is needed for calciﬁcation, the less is available for

other biological processes such as growth or reproduction, reducing the organisms’ weight and overall competitiveness

and viability.

Exposure of external shells to more acidic water can affect their stability by weakening or actually dissolving

carbonate structures. Some of these shells are shielded from direct contact with seawater by a special coating that

the animal makes (as is the case in mussels). The increased energy needed for making the shells to begin with impairs

the ability of organisms to protect and repair their dissolving shells. Presently, more acidic waters brought up from

the deeper ocean to the surface by wind and currents off the Northwest coast of the USA are having this effect on

oysters grown in aquaculture.

Ocean acidiﬁcation affects not only species producing calciﬁed exoskeletons. It affects many more organisms either

directly or indirectly and has the potential to disturb food webs and ﬁsheries. Most organisms that have been

investigated display greater sensitivity at extreme temperatures so, as ocean temperatures change, those species

that are forced to exist at the edges of their thermal ranges will experience stronger effects of acidiﬁcation.

437

Ocean Systems Chapter 6

alcified structures, used for defense and structural support. Calcification

usually occurs in separate body or cell compartments, where pH and thus

–

concentration and saturation Ω (Section 6.1.1) are maintained at

values higher than in other body fluids or ambient water (Taylor et al.,

2011; Trotter et al., 2011; McCullough et al., 2012; Venn et al., 2013).

impedes the formation of carbonate such that calcification rate

decreases. It may be maintained by enhanced transport of ions, incurring

elevated energetic costs (Figure 6-10).

External carbonate structures like shells rely on ambient seawater being

supersaturated with carbonates. Decreasing oceanic carbonate levels

reduce the saturation levels (Ω) of calcite or aragonite in the water.

Reduction to below unity may lead to the corrosion of carbonate shells

(FAQ 6.3). However, many species protect their shells from direct contact

with seawater by various types of organic coating (e.g., a periostracum

in mollusks and brachiopods, an epicuticle covering the carapace of

crustaceans, an epidermis covering the tests of urchins, epithelial tissue

covering aragonite in corals, and coralline algae precipitating CaCO

(mostly Mg-calcite) within their cell wall). A meta-analysis of the effects

f ocean acidification on biological processes indicates that reductions

in the rate of net calcification (calcification minus dissolution) and survival

are the most uniform responses across organisms studied, relative to

other, more variable impacts such as reduced growth, development, and

abundance (Kroeker et al., 2013; see also Box CC-OA).

Some organisms benefit from elevated CO

partial pressures (pCO

Photosynthesis and/or nitrogen fixation in selected microorganisms are

impacted by OA, but effects are species or taxon specific, possibly

depending on how they acquire carbon, that is, the presence and in

particular the type, capacity, and energetic costs of carbon-concentrating

mechanisms (CCMs; Giordano et al., 2005; Kranz et al., 2011).

A comprehensive picture of responses to CO

requires consideration of

variable sensitivities between species and life stages and taxon-specific

sensitivity distributions, as shown by a meta-analysis of animal data

(Wittmann and Pörtner, 2013; see also Figure 6-10b). Echinoderms,

bivalves, gastropods, and corals begin to respond negatively at lower CO

levels than crustaceans or cephalopods (Figure 6-10b). This sensitivity

Taxon

No. of

studies

No. of

parameters

studied

Total no.

of species

studied

pCO

where the most

vulnerable species is negatively

affected or investigated pCO

range

(µatm)

Assessment of

tolerance to

RCP 6.0 (conﬁ dence)

Assessment of

tolerance to

RCP 8.5 (conﬁ dence)

Cyanobacteria 17 5 9+ 180–1250

Beneﬁ cial (low)Beneﬁ cial (low)

Coccolithophores 35 67+ 740Tolerant (low) Vulnerable (medium)

Diatoms 22 528+ 150–1500

Tolerant (low) Tolerant (low)

Dinoﬂ agellates 12 411+ 150–1500

Beneﬁ cial (low) Tolerant (low)

Foraminifers 11 4 22 588 Vulnerable (low) Vulnerable (medium)

Seagrasses 6 6 5 300–21000

Beneﬁ cial (medium)Beneﬁ cial (low)

Macroalgae (non-calcifying) 21 521+

280–20812

Beneﬁ cial (medium)Beneﬁ cial (low)

Macroalgae (calcifying) 38 10 36+ 365Vulnerable (medium) Vulnerable (high)

Warm-water corals 45 13 31 467 Vulnerable (medium) Vulnerable (high)

Cold-water corals 10 13 6445Vulnerable (low) Vulnerable (medium)

Annelids 10 617+ 1200Tolerant (medium) Tolerant (medium)

Echinoderms 54 14 35 510 Vulnerable (medium) Vulnerable (high)

Mollusks (benthic) 72 20 38+ 508Vulnerable (medium) Vulnerable (high)

Mollusks (pelagic) 7 8 8 550 Vulnerable (low) Vulnerable (medium)

Mollusks (cephalopods) 10 8 5 2200 (850 for trace elements) Tolerant (medium) Tolerant (medium)

Bryozoans 738+ 549Tolerant (low) Vulnerable (low)

Crustaceans 47 27 44+ 700Tolerant (medium) Tolerant (low)

Fish

51 16 40 700 Vulnerable (low) Vulnerable (low)

Table 6-3 | Tolerances to ocean acidiﬁ cation in marine taxa, assessed from laboratory and ﬁ eld studies of species in the CO

partial pressure (pCO

) range from <650 to >10000

µatm, compared to present day atmospheric levels of 400 µatm. (It should be noted that anthropogenic CO

emissions add to the natural variability of CO

concentrations

in marine environments, which can reach much higher than atmospheric levels.) Variables studied include growth, survival, calciﬁ cation, metabolic rate, immune response,

development, abundance, behavior, and others. Neither all life stages, nor all variables, including the entire range of CO

concentrations, were studied in all species. Conﬁ dence

is based on the number of studies, the number of species studied, and the agreement of results within one group. + denotes that possibly more species or strains (genetically

distinct populations of the same species) were studied, as only genus or family were speciﬁ ed; beneﬁ cial: most species were positively affected; vulnerable: more than 5% of

species in a group will be negatively affected by 2100; tolerant: more than 95% of species will not be affected by 2100. RCP 6.0: Representative Concentration Pathway (RCP)

with projected atmospheric pCO

= 670 µatm; RCP 8.5: pCO

= 936 µatm in 2100 (Meinshausen et al., 2011). Conﬁ dence is limited by the short- to medium-term nature of

various studies and the lack of sensitivity estimates on evolutionary time scales, that is, across generations (see separate reference list, Online Supplementary Material). Note that

the assessment of variability between species from the same animal phylum has revealed an increase in the fraction of sensitive species with rising CO

levels; see Figure 6-10.

Rather than a sensitivity threshold the entire range of investigated pCO

values is given for groups of photosynthetic organisms. In all studies photosynthetic rates are stimulated

to different, species-speciﬁ c degrees by elevated pCO

, indicating low vulnerability. Coccolithophores and calcifying algae are assessed as being more sensitive than other

photosynthetic organisms due to reduced calciﬁ cation and shell dissolution.

Conﬁ dence levels for ﬁ shes were converted from medium to low, in light of uncertainty on the long-term persistence of behavioral disturbances.

438

Chapter 6 Ocean Systems

−

Ocean

space

orming tissue

]

−

[CO

−

hell/skeleton-

(gills,gut etc.)

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Skin, gills

Respiration rate

Control

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Intracellular

processes

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xchange

Ion-

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rotein

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Metabolic rate

]

−

[HCO

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or muscle heart, brain,.g., eAnimal cell from,

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−

HCO

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−

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(

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occolith

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(c)

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0000

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1–

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370

71–

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500–

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–1370

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–0.60

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ater coralsarm-wWater coralsCold-w

<0.02 >0.02

)

–

Catch rate (tonnes km

439

Ocean Systems Chapter 6

pattern resembles the one seen in the Permian mass extinction (Knoll

et al., 2007; Knoll and Fischer, 2011). The picture for fishes is less clear,

as the present findings of high vulnerability are not met by similar

observations in the fossil record. Evolutionary adaptation may thus

eliminate or minimize reported effects.

The capacity for pH and ion regulation and other relevant processes can

be upregulated by gene expression, as seen in acclimation studies in

echinoderm larvae (O’Donnell et al., 2010; Martin et al., 2011) and fishes

(Deigweiher et al., 2008; Tseng et al., 2013), in warm-water coral branches

(Kaniewska et al., 2012), but not in a study of warm-water coral larvae

(Moya et al., 2012). Few studies address whether and to what extent

species undergo evolutionary adaptation to high pCO

, as seen in the

coccolithophore Emiliania huxleyi over 500 asexual generations

(Lohbeck et al., 2012). In organisms with longer generation times,

perturbation studies in the laboratory measure tolerance and acclimation,

but not adaptation or natural selection. Animal adaptation is accelerated

by high functional variability among larvae, enabling selection of resistant

genotypes (low to medium confidence; Sunday et al., 2011; Parker et

al., 2012; Pespeni et al., 2013). This may explain the selective mortality

seen in Atlantic cod larvae under elevated CO

(Frommel et al., 2012).

Both acclimatization and adaptation will shift sensitivity thresholds but

the capacity and limits of species to acclimatize or adapt remain largely

unknown and hence impacts of acute exposures cannot easily be scaled

up to effects on the longer, evolutionary time scales of ocean acidification

(Wittmann and Pörtner, 2013). Observations in ecosystems characterized

by permanently elevated or fluctuating CO

levels, such as upwelling

areas, OMZs (Section 6.1.1), or seeps, reflect the existence of sensitivity

thresholds (high confidence; Section 6.3.2.5) but organisms may have

evolved a higher resistance to increased CO

levels than elsewhere (low

confidence).

Table 6-3 compiles effects of ocean acidification observed across taxa in

laboratory and field experiments. The latter include studies in mesocosms

and at natural analogs, submarine CO

venting areas at locales such as

Ischia, Italy (Hall-Spencer et al., 2008), Papua New Guinea (Fabricius et

al., 2011), and Puerto Morelos, Mexico (Crook et al., 2012). It should be

noted that anthropogenic CO

accumulation according to RCPs adds to

the natural variability of CO

concentrations in marine environments.

Many groups, especially sessile or non-photosynthetic calcifiers, have

sensitive species with vulnerability thresholds surpassed under RCP6.0

by 2100 (low to medium confidence).

Recent meta-analyses also summarize OA effects, two for biogeochemical

processes and relative effect sizes (Harvey et al., 2013; Kroeker et al.,

2013), one for the distribution of sensitivity between species within

major animal phyla and its change depending on ambient pCO

(Figure

6-10; Wittmann and Pörtner, 2013). All of these analyses consider the

interaction of warming and CO

accumulation (Section 6.3.5). Present

limitations in understanding the mechanisms of effect and their long-

term persistence compounds accurate projections of the long-term

effects of OA (medium confidence; Wittmann and Pörtner, 2013).

6.3.2.2. Microbes

The physiology of both calcifying (coccolithophores) and non-califying

phytoplankton can be influenced by changes in carbonate system

variables caused by ocean acidification (Figure 6-10a). Growth and

photosynthetic rates of diatoms in laboratory cultures are considered

relatively insensitive to elevated CO

(Rost et al., 2003; Trimborn et al.,

2008). Dinoflagellate sensitivity to elevated CO

is poorly studied (Hansen

et al., 2007), but in one species carbon fixation rates were enhanced at

750 µatm CO

while growth remained unaffected (Fu et al., 2008).

Indirect effects of ocean acidification on phytoplankton physiology include

altered availability of trace metals needed for many biochemical cycles

(Hoffmann et al., 2012).

Harmful algal blooms are a growing problem in coastal waters worldwide

(Section 6.4.2.3), and many of the various phytoplankton species that

produce bio-accumulated toxins are sensitive to changes in the seawater

carbonate buffer system (Hallegraeff, 2010; Fu et al., 2012). For example,

the dominance and community structure of harmful bloom dinoflagellates

can be profoundly altered by changing pCO

(Tatters et al., 2013), and

both toxic dinoflagellates and diatoms have been shown to produce

higher toxin levels under near-future levels of ocean acidification (Fu et

al., 2010; Sun et al., 2011).

Some planktonic N

-fixing cyanobacteria (diazotrophs), for example,

strains (genetically distinct populations of the same species) of offshore

cyanobacteria of the genera Trichodesmium and Crocosphaera, respond

to rising CO

with increased rates of both carbon and N

fixation (Fu et

al., 2008; Lomas et al., 2012). In contrast, laboratory studies using the

bloom-forming cyanobacteria Nodularia (an organism largely found in

coastal stratified, eutrophic waters) revealed decreased growth and N

Figure 6-10 | (a) Responses of a schematized marine animal (left) and a phytoplankter (right) to ocean acidiﬁcation. Effects are mediated via diffusive CO

entry (black arrows)

into body and cell compartments, resulting in a rise in pCO

(highlighted in red), a drop in compartmental pH (highlighted in blue), and their effects (red arrows) on various

processes (red text) in tissues and cellular compartments, as well as on calcium carbonate saturation state (Ω) at calciﬁcation sites (after Pörtner, 2008; Taylor et al., 2011).

Variable sensitivity relates to the degree of pH decline and compensation, depending on the capacity of pH and ion regulation. (b) Distribution of sensitivities across species within

animal phyla, under progressively rising water CO

levels, as percent of studied cold- and warm-water coral (mostly scleractinia), echinoderm, molluskan, crustacean, and ﬁsh

species affected negatively, positively, or not at all (for effects considered, see text). As not all life stages, variables, and pCO

ranges were covered in all species, two assumptions

partially compensate for missing data: 1) Negative effects at low pCO

will remain negative at high pCO

. 2) A positive or neutral outcome at both low and high pCO

will be the

same at intermediate pCO

. As responses reported for each species vary for each pCO

range, variable species numbers result (on top of columns). The total number of species

studied in a group is shown as the number above the control column. The control category corresponds to 380 µatm. For 2100, RCP scenarios falling within each CO

partial

pressure (pCO

) category are as follows: RCP4.5 for 500–650 μatm (approximately equivalent to ppm in the atmosphere), RCP6.0 for 651–850 μatm, and RCP8.5 for 851–1370

μatm. By 2150, RCP8.5 falls within the 1371–2900 μatm category. Horizontal lines above columns represent frequency distributions signiﬁcantly different from controls

(Wittmann and Pörtner, 2013). Data for warm- and cold-water corals as in Table 6-3. (c) Areas with reported annual catches of marine calciﬁers (crustaceans and mollusks) ≥

0.005 tonnes km

–2

depicted on a global map (weighted mean of the orange color area =0.07 tonnes km

–2

) showing the distribution of ocean acidiﬁcation in 2100 according to

RCP8.5 (WGI AR5 SPM; pH change from 1986–2005 to 2081–2100) as well as the distribution of warm-water (green dots) and cold-water coral communities (purple dots).

440

Chapter 6 Ocean Systems

ixation under elevated CO

onditions (Czerny et al., 2009). The wide

range of responses in N

fixation (e.g., Hutchins et al., 2007; Levitan et

al., 2007; Kranz et al., 2010) may be explained by different CO

affinities

(i.e., dependences of growth rates on CO

concentration) of a range of

-fixing cyanobacteria (Trichodesmium and Crocosphaera) from different

oceanic biomes. Some species/strains operate at close to maximum

growth rates at present-day oceanic CO

levels, whereas others had

sub-optimal growth rates under these conditions (Hutchins et al., 2013).

To date, the physiological mechanisms underlying these responses remain

unknown, especially in open-ocean nitrogen fixers. Cyanobacteria may

reallocate energy from their energetically expensive CCMs toward N

fixation and the acquisition of growth limiting nutrients (Kranz et al.,

2010; Levitan et al., 2010), but evidence for such diversion of energy is

lacking. Whether nitrogen fixation will increase with progressive ocean

acidification remains to be explored (low confidence, limited in situ

evidence, medium agreement).

The responses of coccolithophore calcification to OA are species specific

and highly variable. The function(s) of calcification are not well understood,

making it difficult to evaluate the consequences of lowered calcification

(e.g., Rost et al., 2008). Reductions, increases, and unchanged calcification

rates (and shell structure) have all been found in different coccolithophore

species for RCP8.5 CO

conditions projected around 2100 (Riebesell et

al., 2000; Zondervan et al., 2001; Langer et al., 2006; Iglesias-Rodriguez

et al., 2008). Calcification in coccolithophores is species (Langer et al.,

2006) and in Emiliania huxleyi even strain specific (Langer et al., 2009,

2011; Hoppe et al., 2011). It thus remains unclear whether OA will result

in exoskeletons that are insufficiently calcified for sustained structural

support and protection in coccolithophores (medium evidence, low

agreement).

Foraminifera display decreasing calcification and shell weight under

elevated CO

(Lombard et al., 2010). Changes in historical specimens

(Moy et al., 2009; see Section 6.3.2.5.1) and during glacial-interglacial

cycles (Barker and Elderfield, 2002) support projections of future reductions

in net calcification by foraminifera (medium to high confidence).

6.3.2.3. Macroalgae and Seagrasses

Primary production, shoot density, reproductive output, and below-

ground biomass of seagrasses generally respond positively to elevated

pCO

, indicating CO

limitation of their productivity. Such effects were

identified in both laboratory and field above 720 to 1800 µatm (high

confidence; e.g., Palacios and Zimmerman, 2007; Hall-Spencer et al., 2008;

Andersson et al., 2011; cf. Section 5.4.2.3). Production, growth, and

recruitment of most but not all non-calcifying seaweeds also increased

at CO

levels from 700 to 900 µatm (RCP8.5; Porzio et al., 2011; Kroeker

et al., 2013). Some non-calcifying seaweeds and seagrasses will thus

benefit from future ocean acidification (high confidence) but OA exposes

them to higher than usual grazing as a consequence of losing deterrent

phenolic substances (low confidence; Arnold et al., 2012).

Calcifying algae (corallines) show complex and species-specific responses

of photosynthesis to elevated CO

, but calcification is impacted once

species-specific pCO

thresholds are surpassed (medium confidence;

Anthony et al., 2008; Martin and Gattuso, 2009). At habitat temperature

alcification by temperate coralline red and calcareous green algae

increased at CO

levels up to 900 µatm and decreased only at the highest

concentration applied (2850 µatm), but did not fall below rates found

at present-day pCO

(Ries et al., 2009). During 3 months of exposure,

growth of Lithothamnion glaciale, a cold-water calcareous red alga,

decreased progressively with rising CO

levels, and its structural integrity

was weakened beyond 590 µatm (Ragazzola et al., 2012), potentially

influencing ecosystem function. Some calcifying algae may thus be

impacted by future ocean acidification (medium confidence).

6.3.2.4. Animals

Studies of marine animals and their life stages show a high diversity

and variability of processes affected by ocean acidification. Many

variables studied reflect physiological performance (O

consumption,

exercise, behavior, calcification, growth, immune response, acid-base

balance, gene expression, fertilization, sperm motility, developmental

time, production of viable offspring, and morphology; Table 6-3; Figure

6-10). In some species growth may be stimulated by OA, in others

depressed or unaffected (cf. Gooding et al., 2009; Munday et al., 2009a,

2011a; Dupont et al., 2010). The degree of CO

-induced acidosis and its

compensation by ion exchange may shape sensitivity (Section 6.3.2.1).

Full exploitation of the ability to resist pCO

increases depends on the

availability and high quality of food and the strengthening of fitness

(Gooding et al., 2009; Melzner et al., 2011). However, food quality of

prey organisms may decrease under elevated pCO

. For example, slower

reproduction and growth of the copepod Acartia tonsa under 760 µatm

pCO

was related to the decreasing quality of its diatom food (Rossoll

et al., 2012).

Changes in calcification rates reported from CO

manipulation

experiments vary widely. Reduced calcification and weakened calcified

structures were seen under elevated pCO

in corals (see Section 6.3.2.4.2),

echinoderms (Kurihara and Shirayama, 2004), mollusks (Gazeau et al.,

2013), and larval crustaceans (Arnold et al., 2009; Walther et al., 2011).

Some adult limpets and urchins increased calcification rates at pCO

from 600 to 900 µatm, before it fell at even higher pCO

. In some adult

crabs, lobsters, and shrimps calcification rates increased further with

rising pCO

(Ries et al., 2009). Stronger internal structures such as

cuttlebones and otoliths resulted from enhanced calcification under

elevated pCO

in juvenile cuttlefish (cephalopods: Gutowska et al.,

2008) and fishes (Checkley, Jr. et al., 2009; Munday et al., 2011b), with

unclear impacts on fitness. Energy costs in epithelia or calcification

compartments may be enhanced by elevated pCO

causing a stimulation

of metabolism (Section 6.3.2.1). In some cases, this may indicate

imbalances in energy budget rather than increased CO

resistance, for

example, if costs are down-regulated in muscle or liver. Enhanced

calcification can then occur at the expense of growth (medium confidence;

Wood et al., 2008; Beniash et al., 2010; Thomsen and Melzner, 2010;

Parker et al., 2011).

Studies on calcifying zooplankton focused on pteropods (planktonic

mollusks with aragonite shells). These form an integral part of the food

web, both as grazers and prey, for example, for pink salmon (Armstrong

et al., 2005; Hunt et al., 2008). In the Sub-Arctic, the Arctic, and the

Southern Ocean, pteropods will reduce calcification in response to OA

441

Ocean Systems Chapter 6

ntil at least the end of the century (medium confidence; Orr et al., 2005;

Comeau et al., 2009; Lischka et al., 2011).

Elevated CO

causes behavioral disturbances in fishes (studied mostly

in larvae and juveniles; Munday et al., 2010; Ferrari et al., 2011;

Domenici et al., 2012; Jutfeld et al., 2013) through neural mechanisms

(Nilsson et al., 2012). The long-term persistence and evolutionary

relevance of these behavioral effects need further study before general

conclusions can be drawn (low confidence; Wittmann and Pörtner, 2013;

see also Section 6.3.5).

6.3.2.5.3. Macrophytes and macrofauna

Macrofauna and macrophyte communities have been studied in

mesocosms and in ecosystems exposed to shifted upwelling regimes or

at natural volcanic CO

vents (Fabricius et al., 2011; Kroeker et al., 2011).

The latter are considered as natural analogs of future ocean acidification.

An 8-year trend of (variable) pH decline in upwelled waters along the

Northeast Pacific coast was paralleled by shifts in community composition,

where shelled species like mussels were replaced by fleshy algae and

barnacles (Wootton et al., 2008). Macrofaunal calcifiers at CO

vents

(Hall-Spencer et al., 2008; Fabricius et al., 2011) and in mesocoms

(Christen et al., 2013) display a lowering of species richness. These

findings suggest that non-calcifiers increasingly outcompete calcifiers

once pH

decreases to a mean of 7.8 to 7.7 (medium confidence).

Finally, a loss of calcifiers from mesocosms occurred around 0.5 units

below the pH values expected from OA under RCP8.5 by 2100 (medium

confidence; Christen et al., 2013). At CO

seeps, calcitic bryozoans replace

coralline algae, which have more soluble high-calcite skeletons (Martin

et al., 2008). Seagrasses and non-calcifying algae gain a competitive

advantage (Fabricius et al., 2011). Coral communities exposed to high

pCO

waters (from upwelling or seeps) have lower growth, calcification,

and biodiversity (Manzello et al., 2008; Fabricius et al., 2011), resulting

in a shift from net accretion to erosion (Box CC-CR). The use of seeps

as analogs of future OA is limited as pH variability is high at these sites,

such that effective values may be lower than indicated by the average

change (Hall-Spencer et al., 2008; Porzio et al., 2011). During periods

of high pH at the seeps, they are recolonized by invertebrates and

fishes from neighboring areas with normal pH, compromising

assessments of long-term sensitivity thresholds. Overall, findings

available from mesocosms and natural analogs indicate losses in

diversity, biomass, and trophic complexity of benthic marine

communities due to elevated CO

(high confidence) and support the

projection of similar shifts in other systems with continued OA

(medium confidence).

Enhanced freshwater input by poorly buffered rivers or by precipitation,

into estuaries, brackish oceans like the Baltic (Section 30.5.3.1.4), and

into freshening polar oceans, reduces salinity and alkalinity at rising

atmospheric pCO

and thereby, alters the carbonate system and enhances

OA (Section 6.1.1). Estuaries usually have OMZs, where background

pCO

is elevated. Its reduction by dilution causes the acidification effect

to be somewhat less. Enhanced pH reduction and variability in

hyposaline waters may constrain the distribution of sensitive species

further (low confidence; Miller et al., 2009; Denman et al., 2011).

443

Ocean Systems Chapter 6

6.3.2.5.4. Conclusions

Natural analogs and laboratory and mesocosm experiments provide

evidence for differential effects of ocean acidification on species and

communities. Sensitivity to OA is species specific (high confidence);

differential sensitivities and associated shifts in performance and

distribution will change predator-prey relationships and competitive

interactions (low to medium confidence). OA may stimulate global net

primary production (low confidence) and nitrogen fixation (medium

confidence). OA will increase the abundance and primary production of

non-calcifying macrophytes, but will be harmful to calcifying algae and

many heterotrophs (medium confidence). Ecosystems relying on calcified

structures and at risk of dissolution under OA include warm-water coral

reefs (high confidence) and their cold-water equivalent (medium

confidence). Further studies need to explore how OA may change the

composition of communities, impact food webs, and affect higher

trophic levels.

6.3.3. Life in Hypoxia and Anoxia

6.3.3.1. Principles

Hypoxia constrains organisms which rely on aerobic metabolism

(Section 6.1.1; FAQ 6.2). Below O

concentrations of 60 µmol kg

–1

commonly termed hypoxic (Section 6.1.1.3), communities undergo

species losses and replacements and are transformed into communities

with species showing characteristic hypoxia adaptations. However, O

can limit animal life at even higher levels, just below air saturation (Gilly

et al., 2013). Organisms’ tolerance thresholds have been defined by

either the critical O

partial pressure (P

) or concentration (O

crit).

Thresholds vary across domains and are highest for large multicellular

organisms. Among these, the P

at rest varies depending on species,

body size, and life stage. In animals below the P

aerobic metabolic rate

fails to be maintained and anaerobic metabolism contributes to energy

production (Pörtner and Grieshaber, 1993). The critical oxygen threshold

is set by the capacity of ventilatory and circulatory systems to supply

and cover demand. The threshold increases once metabolism is

stimulated by muscular activity, temperature, or food uptake (Pörtner,

2002a; Ekau et al., 2010; Seibel, 2011; see also Figure 6-11). At extreme

temperatures, O

crit approaches the oxygen content of air-saturated

water (Pörtner, 2010; McBryan et al., 2013), indicating high sensitivity

to hypoxia in the warmth. Most animals can only sustain anaerobic

metabolism temporarily, even if they are energy efficient and survive

long periods of anoxia (Grieshaber et al., 1994). Such time-limited

tolerance is higher in large than in small individuals or larvae, related

to the higher capacity of anaerobic metabolism in large specimens (Gray

et al., 2002; Jessen et al., 2009).

6.3.3.2. Microbes

Bacteria and protists consume ambient oxygen down to very low

levels in oxygen minimum zones and sustain OMZs by their metabolic

diversity (Figure 6-11; WGI AR5 Section 3.8.3). OMZs form habitat for

both anaerobic and aerobic microbes that can utilize very low (<1 µmol

–1

) O

concentrations (Stolper et al., 2010). Hypoxia is paralleled by

levated pCO

nd enhanced acidification. Expanding OMZs will select

for the proliferation of spezialized microbes (high confidence).

6.3.3.3. Animals and Plants

In mesopelagic OMZs, zooplankton also contribute to the development

of hypoxia (Robinson et al., 2010; see also FAQ 6.2). During daytime

zooplankton congregate at the upper margin of OMZs, where the

degradation of organic material causes intensified respiration and oxygen

depletion (Bianchi et al., 2013). Animals living permanently in the OMZ

still cover virtually all energy demand by aerobic metabolism. This requires

special adaptations leading to a reduction in O

and energy demand,

and the improved ability to use available O

efficiently. Enhanced

hypoxia tolerance reflected in low O

crit values is supported by small body

size and by cold temperature (Vetter et al., 1994; Pörtner, 2002b; Levin

et al., 2009). Accordingly, low O

levels support abundant meiofauna

(very small fauna, <1 mm) that benefit from abundant food and reduced

predation by larger organisms (Levin, 2003). Under suboxia only

specialists can survive (Vaquer-Sunyer and Duarte, 2008). Expansion of

suboxic and anoxic centres of pelagic OMZs and benthic dead zones

will lead to loss of habitat for animal life (high confidence).

Large, more active animals such as fishes, crustaceans, and muscular

(as opposed to ammoniacal) squids tend to have high O

demands

associated with high O

crit thresholds, and are therefore excluded from

permanently hypoxic water bodies. However, even in high-activity animal

groups some specialists such as Humboldt squid or bigeye tuna have

adapted to enter hypoxic environments though only temporarily (Richards

et al., 2009; Seibel, 2011). The time-limited tolerance of animals to

hypoxia below the O

crit is maximized by the depression of energy demand,

for example, during periods of metabolic arrest (e.g., developmental

arrest or diapause of copepods; Auel et al., 2005). Hypoxia-adapted

lifeforms will benefit from expanding OMZs (high confidence).

There is little information on the hypoxia sensitivity of macrophytes or

their O

crit values. In eelgrass (Zostera marina), warming causes the

hypoxia threshold to rise due to a strong increase in tissue respiration.

Concomitant water or sediment hypoxia can elicit tissue anoxia and

sudden die-offs (Raun and Borum, 2013). By contrast, macroalgae

attached to rocks rarely encounter anoxia (Raven and Scrimgeour, 1997).

Expanding benthic OMZs will constrain the distribution of macrophytes

(medium confidence).

6.3.3.4. Ecosystems

OMZs, shoaling, and expanding vertically and laterally (Gilly et al., 2013)

will cause habitat and abundance losses for intolerant taxa such as

mesopelagic (Koslow et al., 2011) and epipelagic fishes with a high O

demand (medium confidence; Prince et al., 2010; Stramma et al., 2012;

see also FAQ 6.2). In line with the distribution of hypoxia sensitivities

(Figure 6-11; Sections 6.3.3.1, 6.3.3.3), expanding OMZs will further

constrain the distribution of key zooplankton and nekton species and

influence their diurnal and ontogenetic vertical migrations (medium

confidence; Ekau et al., 2010). The composition of microbial and faunal

pelagic communities will shift from diverse mid-water assemblages to

444

Chapter 6 Ocean Systems

igrant biota that return to oxygenated surface waters at night (Seibel,

2011). Dissolved O

, among other factors, plays an important role in

shaping large alternating fluctuations of sardine and anchovy abundances,

particularly off Peru. Anchovies are not strongly affected by a shallow

oxycline (<10 m), while sardines actively avoid such conditions

(Bertrand et al., 2010). Where OMZs intersect the continential shelves,

groundfishes (McClatchie et al., 2010) and large benthic invertebrates

such as crabs display high mortalities (Chan et al., 2008). Susceptibility

of early life stages to hypoxia in both pelagic and benthic ecosystems

(

Ekau et al., 2010) threatens population survival. Effects of hypoxia

propagate along the food chain, constraining fish stocks and top

predators (high confidence; Stramma et al., 2010). Hypoxia reduces

biodiversity (Levin et al., 2009; Gooday et al., 2010) and causes the

marginalization of calcifiers, due to low metabolic rates and high pCO

(high confidence; Levin, 2003; Levin et al., 2009).

The expansion and enhanced variability of OMZs increases dissimilatory

nitrate reduction and anaerobic ammonium oxidation (anammox), both

releasing N

into the atmosphere, reducing the availability of fixed

nitrogen, and limiting oceanic primary productivity (medium confidence).

Water column denitrification and N

fixation are spatially and temporally

variable (limited evidence, low confidence), suggesting that climate

effects on these processes are unlikely to operate uniformly (Brandes

et al., 2007; Fernandez et al., 2011; Franz et al., 2012).

If O

levels decline and OMZs expand, tolerant taxa, such as anaerobic

bacteria (Ulloa et al., 2012), gelatinous zooplankton (medusae,

ctenophores), selected fishes (gobies, hake), and possibly selected

cephalopods (Gilly et al., 2006; Bazzino et al., 2010) will respond with

range expansions or population growth. Similar phenomena are

expected with intensified upwelling causing extensive mortalities of

coastal fishes and invertebrates (Box CC-UP). A community change

toward hypoxia-tolerant fauna will occur in mid-water (high confidence).

The diversity of macroorganisms will decrease and, finally, higher

marine organisms will disappear and heterotrophic microorganisms will

dominate (high confidence). In isolated water bodies such as the Black

Sea, warming will lead to the expansion of anoxia and hydrogen sulphide

S) poisoning, reduce pelagic and bottom faunal distributions, and

shape trophic relations, energy flows, and productivity (Daskalov, 2003;

Fashchuk, 2011).

6.3.4. Mixed Layer Depth and

Light Shaping Net Primary Production

The upper ocean is characterized by physical and chemical gradients in

the surface mixed layer that influence the magnitude of photosynthetic

carbon fixation, often termed net primary production (NPP). The availability

of light and nutrients to photoautotrophs sets daily rates of NPP and

may be altered directly or indirectly, through changing mixed layer

depths, shifts in the circulation regime at different spatial scales, and

the physical displacement of organisms (Section 6.1.1.4; Box CC-PP;

Figure 6-2). A changing climate will affect mixed layer depth, cloudiness,

and/or sea ice areal extent and thickness and thereby modulate NPP

(high confidence). A stronger vertical density gradient will reduce the

communication between the sunlit upper ocean where photosynthesis

takes place and the underlying nutrient-rich waters (Figure 6-2). The

supplies of plant nutrients (macro-nutrients) such as nitrate, and of

micro-nutrients such as iron (Pitchford and Brindley, 1999) vary seasonally

(Boyd, 2002) and regionally (Moore et al., 2002), such that NPP may be

simultaneously limited (co-limited) by more than one resource (Saito et

al., 2008; see also Section 6.3.5).

The changing range and intensity of underwater light will lead to

changes in NPP as well as in phytoplankton community composition

(Doney, 2006; Boyd et al., 2010). The response of phytoplankton to

“

”

OMZ = Oxygen

minimum zone

POM = Particulate

rganic matter

“Dead Zone” = Anoxic

+ Sub-oxic zone

hytoplankton

bloom

nhanced

stratiﬁcation

idwater OMZ

espiration

OMZ formation

pwelling

urface ocean

POM

armful algal

bloom

Nutrient input

Upwelling

inds

Acidiﬁcation

Estuary

oastal ocean

Open ocean

N-fertilization

ight and

arming

Hypoxia-adapted

communities

Hypoxia-sensitive communities

Oxic

Present

Projected

OMZ

Environmental O

concentration high low

Large multicellular Eukarya

Small multicellular Eukarya

Specialized unicellular Eukarya, Bacteria, and Archa ea

Figure 6-11 | (a) Principal mechanisms underlying the formation of hypoxic conditions

and their biological background (modiﬁed from Levin et al., 2009; Levin and Sibuet,

2012). The buoyancy ﬂux from ﬂuvial discharges produces sharp density stratiﬁcation

at the base of the freshened layer (also valid for ice melt and high precipitation) near

the surface and, hence, vertical mixing is greatly reduced. In consequence, the nutrient

inputs from the river and the atmosphere accumulate in a narrow upper layer, leading

to blooms of phytoplankton, possibly including harmful algae. The enhancement of

oxygen consumption due to aerobic decomposition of sinking particulate organic

matter (POM) results in hypoxic conditions of benthic and mid-water oxygen minimum

zones (OMZs). Enrichment of nutrients (eutrophication) results in coastal dead zones.

In the open oceans, heating of the upper layer increases stratiﬁcation, while the

wind-driven upwelling of hypoxic, nutrient-rich water from deeper layers adds to the

formation of the OMZs (Box CC-UP). (b) Distribution of free-living marine organisms

(microbes such as archaea, bacteria, protists, small and large multicellular animals, and

plants) across the ranges of O

concentrations in various water layers. Hypoxia

tolerance is enhanced in small compared to large organisms, allowing unicellular

species and small animals to thrive in extremely hypoxic habitats. Species richness and

body size of animals decrease with falling O

levels.

445

Ocean Systems Chapter 6

hanging sunlight involves photo-physiological acclimation via changes

in cellular chlorophyll, but such acclimation is constrained by unidentified

limits (Falkowski and Raven, 1997). A longer growing season, with more

sea ice-free days between 1998 and 2009, may have increased NPP in

open Arctic waters (Arrigo and van Dijken, 2011; see also Box CC-PP),

complemented by massive under-ice blooms as seen in 2011, favored

by light that penetrates surface melt ponds and thinner, for example,

first-year ice (Arrigo et al., 2012). There are also reports of increased

incidences of high phytoplankton stocks, and hence of greater NPP,

deeper in the water column (i.e., where it cannot be detected by satellite)

during summer in the Arctic, which have implications to assessing

changes in NPP from space (Hill et al., 2013). Little is known about shifts

from sea ice algae to free-drifting phytoplankton expected with a

decrease in sea ice cover and effects of increased light in polar waters

in the coming decades (low confidence). In the Arctic, summer ice melt

led to a rapid export of sea-ice algae to the deep ocean (Boetius et al.,

2013). As some krill feed primarily on sea ice algae, it is unclear (low

confidence) whether they will adapt to feeding mainly on free-drifting

phytoplankton (Smetacek and Nichol, 2005).

A range of time series observations, from in situ phytoplankton

abundances to satellite remote sensing, have been used to assess

whether phytoplankton stocks and hence rates of NPP have altered over

recent decades. Increases in phytoplankton stocks were found in regions

where colder waters had warmed in the Northeast Atlantic, whereas

the opposite trend was observed for warm-water regions from a

phytoplankton abundance time series (Richardson and Schoeman,

2004). Lower chlorophyll concentrations at warmer SSTs in nutrient-

poor low-latitude waters, based on satellite ocean color data, have been

interpreted as an effect of increased stratification on phytoplankton stocks.

It has thus been suggested that expanding, permanently stratified, low-

chlorophyll, tropical regions (WGI AR5 Chapter 3) indicate declining

phytoplankton stocks in the warming oligotrophic waters of the North

and South Pacific and North and South Atlantic (limited evidence, low

agreement due to methodological uncertainties; Box CC-PP; Polovina

et al., 2008; Signorini and McClain, 2012; see also Section 30.5.1.1.2).

Furthermore, a transition to conditions favoring increased frequency or

even permanence of El Niño in a warmer future (Wara et al., 2005) and

further expansion of subtropical ocean gyres (Polovina et al., 2008;

see also Section 30.5.6) may lead to lower global ocean NPP (low to

medium confidence).

However, these long-term “blended” projections (i.e., constructing a

biomass time series using multiple proxies such as ocean transparency)

of a global decrease in phytoplankton biomass (Boyce et al., 2010) have

been refuted (Mackas, 2011; McQuatters-Gollop et al., 2011;

Rykaczewski and Dunne, 2011). Time series shorter than 20 years do

not resolve impacts of bi-decadal variation such as the Pacific Decadal

Oscillation or the lunar nodal cycle (e.g., Watanabe et al., 2008; Henson

et al., 2010). Analysis of continental shelf ecosystems, including field

data in the most productive upwelling areas covering the last 20 years

(e.g., Chavez et al., 2011), revealed a large variety of trends at scales of

several decades but a general increase in NPP on most shelves (Sherman

and Hempel, 2009; Bode et al., 2011), possibly caused by natural climate

variability, anthropogenic climate change, and/or anthropogenic

eutrophication. Recent field measurements document increasing quantities

of both anthropogenic fixed N (Duce et al., 2008) and biologically fixed

tmospheric nitrogen (Mouriño-Carballido et al., 2011) entering the

open ocean, which could lead to increased NPP especially in warm,

stratified tropical and subtropical oceans provided sufficient phosphate

and other growth requirements are present (low confidence; e.g., Sohm

et al., 2011).

For heterotrophs, from bacteria to fish, mammals, and birds, the uptake

of organic material as food, ultimately provided by NPP, is central not

only to productivity but also for fueling energy-consuming functions

including the resistance of organisms to environmental change and

pathogens (Sections 6.3.1-2). Any direct influence of climate on the

abundance and quality of feed organisms will thus translate to indirect

effects on the productivity and well-being of foraging animals (high

confidence; Figures 6-5a, 6-7a, 6-12).

Overall, pelagic systems respond to climate change by region-specific

changes in productivity with the projection of a small net reduction in

global ocean NPP by 2100 (medium confidence; Box CC-PP). The spatial

reorganization of NPP between latitudes affects higher trophic levels

by alteration of the composition and functioning of pelagic communities

(medium confidence).

6.3.5. Concurrent Responses to Multiple Drivers

Climate change alters oceanic properties globally, with concurrent

changes in temperature, dissolved CO

and O

, light, and nutrient

concentrations (e.g., Sarmiento et al., 1998; Matear and Hirst, 1999;

Boyd and Doney, 2002; Ekau et al., 2010; see also Figure 6-2). Additional

direct human interventions at regional scale comprise the introduction

of non-native species, overfishing, pollution, long-range atmospheric

transport of nitrogen, point-source eutrophication, and habitat destruction

(Carlton, 2000; Boyd and Hutchins, 2012). Worldwide alterations in

marine ecosystems (Pauly et al., 1998; Österblom et al., 2007) have been

linked to direct human activities, especially fishing (Frank et al., 2005;

deYoung et al., 2008; Casini et al., 2009), but may also be caused to

some extent by climate variability and change (Cheung et al., 2013a).

Alteration of each individual property has pronounced effects on

organisms from microbes to animals, and hence on ecosystems

(Sections 6.3.1-4). The cumulative effects of these factors will result in

complex patterns of change, from organismal physiology to the areal

extent and boundaries of biogeographic regions (Table 6-4). In many

organisms, effects of ocean acidification interact with those of other

key drivers such as temperature and hypoxia (Boyd, 2011; Gruber, 2011;

Pörtner, 2012) and translate from molecular to ecosystem level impacts.

In phytoplankton, low light (Zondervan et al., 2002) or nitrogen limitation

(Sciandra et al., 2003) limit beneficial OA effects on photosynthesis and

have a strong negative effect on plankton calcification (Rokitta and Rost,

2012). Nutrients and light support functional adjustments to OA through

gene expression changes (Dyhrman et al., 2006; Richier et al., 2009).

Similar to today, paleo-events such as the Palaeocene-Eocene Boundary

demonstrate concurrent warming, enhanced stratification of the oceans,

deoxygenation of deeper waters, and OA, albeit at a rate more than 10

times slower than today’s rate (Section 6.1.2). Both the complexity of

paleo-ecosystem changes and the complexity of present effects confound

446

Chapter 6 Ocean Systems

he clear attribution of biological trends to individual drivers (Parmesan

et al., 2011). For warming and hypoxia, changes are accelerated by

effects of shifting seasonal or even diurnal extremes and their frequency

on organisms and ecosystems (medium evidence, medium agreement)

(e.g., Pörtner and Knust, 2007; Díaz and Rosenberg, 2008). This may also

apply to effects of anthropogenic OA (limited evidence, low agreement).

6.3.5.1. Principles

Effects of various climate drivers on ocean ecosystems are intertwined

and effects may be exacerbated by responses of biota. For example,

warming reduces O

solubility and enhances biotic O

demand, which

exacerbates hypoxia, produces CO

, and causes acidification (Millero, 1995;

Brewer and Peltzer, 2009). Drivers act with either additive, synergistic (i.e.,

amplification of) or antagonistic (i.e., diminution of) effects. A meta-

analysis of 171 experimental studies that exposed marine systems to two

or more drivers identified cumulative effects that were additive (26%),

synergistic (36%), or antagonistic (38%) (Crain et al., 2008). Effects range

from direct impacts of ocean warming on organismal physiology (Pörtner

and Knust, 2007) to ocean acidification acting together with warming,

for example, on coccolithophore calcite production and abundances

(Feng et al., 2009), or with hypoxia and/or salinity changes (Table 6-4).

Interactions of predominantly temperature, ocean acidification, and

hypoxia have likely been involved in climate-driven evolutionary crises

during Earth history (Pörtner et al., 2005; see also Section 6.1.2).

ffects on individual organisms may also reflect intertwined impacts of

ocean warming, acidification, and hypoxia, which may operate through

interrelated functional principles (Pörtner, 2012). Such knowledge helps

to reconcile apparently contrasting findings. For example, warming

toward the thermal optimum (Figure 6-5a) stimulates resistance to OA;

-induced disturbances of growth and calcification were reversed by

concomitant warming (Findlay et al., 2010; Sheppard-Brennand et al.,

2010; Walther et al., 2011). Warming to above optimum temperatures,

however, constrains performance and exacerbates sensitivity to hypoxia

and/or elevated CO

(Figure 6-5, e.g., via decreased calcification;

Rodolfo-Metalpa et al., 2011). Both hypoxia and/or elevated CO

in turn

enhance heat sensitivity, as seen for CO

in crustaceans (via decreased

heat limits: Walther et al., 2009; Findlay et al., 2010), coral reef fishes

(via reduced performance: Munday et al., 2009b), and corals (via

decreased calcification and CO

-enhanced bleaching: Reynaud et al.,

2003; Anthony et al., 2008). This translates into a narrowing of the

thermal niche (Walther et al., 2009; see also Figure 6-5), which will shrink

biogeographic ranges, affect species interactions, and shift phenologies

(Figure 6-7a). Hence, extreme warming and hypoxia exacerbate CO

effects and vice versa (medium confidence). Such principles need to be

reconfirmed across organism taxa (Pörtner, 2012).

Differences in organism adaptation to a climate zone’s characteristic

temperatures, temperature variability, oxygen content, and ocean

chemistry may shape vulnerability to climate change. In high polar

species evolutionary cold adaptation enhances vulnerability to warming

Biological organization

studied at ecosystem

level

Anthropogenic forcing

Single environmental driver

Multiple environmental

drivers

Fishing / foodwebs Fishing / climate change

Individuals

Lab experiments and field

observations show that warming

alters organismal physiology

and thereby growth (Pörtner and

Knust, 2007).

Shipboard manipulation

experiment addressing interactive

effects of temperature and CO

on coccolithophore calcification

(Feng et al., 2009).

NA Unknown

Population

Physiological effects of warming

change population abundance in

situ (Pörtner and Knust, 2007).

Lab cultures show how altered

pH elicits different responses of

coccolithophore species (Langer

et al., 2006).

Lab cultures show differential

responses of cyanobacterial

groups to temperature and CO

(Fu et al., 2007).

Altered maturation age and

growth rate of populations due to

fishing (Fairweather et al., 2006;

Hseih et al., 2006).

Interactive effects on cod

populations of fishing and

alteration of salinity (Lindegren et

al., 2010).

Ecosystem

Mesocosm experiments

simulating the effect of individual

drivers (e.g., ocean acidification

effects on benthos: Christen

et al., 2013; and on pelagic

communities: Riebesell et al.,

2013).

Mesocosm experiments studying

differential effects of light and

temperature, on copepods versus

diatoms (Lewandowska and

Sommer, 2010).

Effects of fishing on ecosystem

structure — trophic cascades

(Frank et al., 2005).

Interplay of fishing and climate

pressures on ecosystems

promotes lower trophic levels

(Kirby et al., 2009);

enhances diversity loss in benthic

communities (Griffith et al.,

2011).

Biome

Time series observations on

warming and geographical

shifts of zooplankton biomes

(Beaugrand et al., 2009).

Unknown Unknown Unknown

Table 6-4 | Potential interactions between modes of anthropogenic forcing (environmental; foodwebs; harvesting) on different levels of biological organisation. These

interactions, from simple to complex, are illustrated with examples from the published literature. Unknown denotes no published information is available for each of these

categories. NA denotes not applicable for this category.

Approaches: = Experiments (lab or ﬁ eld) = Observations = Modeling = Not applied

447

Ocean Systems Chapter 6

(

medium confidence). In OMZs, marine sediments, and in polar waters

(due to high solubility in the cold), CO

levels are elevated and adaptation

may reduce sensitivity and reliance on calcified structures (Clark et al.,

2009; Walther et al., 2011; Maas et al., 2012). The observed shift from

“overcalcified” to “weakly calcified” coccolithophores Emiliania huxleyi

in cold waters may reflect a related shift in ecotype dominance (limited

evidence, medium agreement; Cubillos et al., 2007).

Despite such potential adaptation, polar calcifiers exposed to higher

and lower carbonate saturation levels have been hypothesized to

be highly sensitive to further CO

accumulation (limited evidence, high

agreement; Orr et al., 2005). Here it appears relevant that cold temperature

reduces energy demand and thereby lowers resistance to ocean

acidification. Both energy demand and resistance are higher in eurytherms

than in high polar and deep sea stenotherms (limited evidence, medium

agreement; Pörtner, 2006; e.g., crustaceans: Pane and Barry, 2007: cf.

Whiteley, 2011). In turn, tropical species may be more sensitive than

temperate zone species (Pörtner et al., 2011). This rough differentiation

of sensitivity is complicated by the local adaptation of populations from

within-species genetic variability (low confidence).

Temperature influences hypoxia sensitivity (Section 6.3.3). Warming

causes the minimum tolerated O

level to rise, enhancing vulnerability

(high confidence). Conversely, hypoxia enhances vulnerability to warming

in animals. This may occur fastest in warm oceans, where metabolic

rates are higher and animals live closer to upper thermal limits (medium

confidence; Pörtner, 2010). However, evolutionary adaptation has led to

high hypoxia tolerance (low P

or O

crit values) in some warm-adapted

coral reef fishes. Further warming then causes a rise in P

which cannot

be compensated for (Nilsson et al., 2010). Limits to hypoxia adaptation

coincide with upper thermal limits (medium confidence).

Complexity in responses rises with the number of drivers involved.

Enhanced river runoff and increased precipitation cause a shift from

marine to more brackish and even freshwater communities, with unclear

consequences for effects of other drivers. Falling primary production

reduces resilience of higher trophic levels (Kirby and Beaugrand, 2009;

Stock et al., 2011). The introduction of non-indigenous species, when

supported by climate-induced shifts in interactions, may promote the

displacement of ecotypes and shifts in ecosystem functioning, for

example, in the Mediterranean Sea (Occhipinti-Ambrogi, 2007; Coll et

al., 2010).

6.3.5.2. Microbes

Both synergistic and antagonistic effects of multiple drivers on microbial

biota in the surface ocean have been observed in manipulation or

modeling experiments (Folt et al., 1999; Boyd et al., 2010; Gruber, 2011).

The productivity of many microbes was simultaneously limited by, for

example, availability of nitrate and phosphate, cobalt and iron (Saito et

al., 2002; Bertrand et al., 2007), or iron and light (Boyd et al., 2010; see

also Section 6.2.2). Warming and high CO

synergistically enhanced

photo-physiological rates of the cyanobacterium Synechococcus,

whereas the cyanobacterial group Prochlorococcus showed no change

(Fu et al., 2007). The magnitude of CO

effects on growth, fixation rates,

or elemental ratios within single species is often strongly modulated by

utrient availability and light conditions (e.g., Sciandra et al., 2003;

Zondervan et al., 2002; Kranz et al., 2010). Such differences cause floristic

shifts in phytoplankton with the potential to restructure predator-prey

interactions (Table 6-4).

Co-limiting factors vary by group, such as nitrogen fixers (e.g., Hutchins

et al., 2007; Kranz et al., 2010), diatoms (Boyd et al., 2010), and

coccolithophores (e.g., Feng et al., 2009; Rokitta and Rost, 2012). This

limits the ability to project climate change effects (Boyd et al., 2010).

The most reliable projections at ocean basin scale come from modeling,

which mainly points to synergistic effects, such as those of elevated

, hypoxia, and warming. For example, OA is projected to alter sinking

particles (C:N ratio and/or reduced calcite content and slower sinking)

with a consequent knock-on effect on water column O

demand already

stimulated by warming, thereby causing expansion of OMZs (Gruber,

2011).

6.3.5.3. Animals and Plants

High oxygen availability alleviates thermal stress as seen in fish and

mollusks (Mark et al., 2002; Pörtner et al., 2006). Conversely, hypoxia

reduces heat tolerance (Section 6.3.5.1), but acclimation to hypoxia

compensates for this and increases thermal tolerance (Burleson and Silva,

2011), for example, by enhancing blood pigment content or reducing

energy demand. Tolerances to hypoxia and to high temperature may

positively correlate in some fishes, indicating potential for adaptive

evolution under climate change (low confidence; McBryan et al.,

2013).

As a consequence of hypoxia narrowing thermal ranges (Section 6.3.5.1),

combined warming and expanding hypoxia may cause mid-water

mesopelagic and demersal fish stocks to decline at rates much quicker

than anticipated in the California Current Ecosystem (McClatchie et al.,

2010; Koslow et al., 2011). In benthic fauna, warming will also increase

vulnerability to hypoxia. Experiments showed a rise in lethal oxygen

concentrations by 25% and thereby reducing survival by 36% at 4°C

warmer temperatures (Vaquer-Sunyer and Duarte, 2011). Hence, warming

is expected to expand the area of ecosystems affected by hypoxia even

if oxygen concentrations remain unchanged (high confidence). Under

combined hypoxia and warming, CO

can extend short-term passive

tolerance (despite constraining long-term tolerance). It facilitates a

reduction in energy demand (Reipschläger et al., 1997; Pörtner et al.,

2000), thereby extending survival of transient extremes of temperatures

or hypoxia (medium confidence).

In macroalgae (non-calcifying) light availability modulates the response

to elevated pCO

and temperature levels (Russell et al., 2011; Sarker et

al., 2013). In warm-water corals, warming acting synergistically with

reduces calcification and increases sensitivity to bleaching (high

confidence; Anthony et al., 2008). Combined warming and OA following

SRES B1 (≈RCP4.5, reduced emission) and A1FI (≈RCP8.5, business-as-

usual) scenarios in mesocosms caused losses of symbionts and corals,

and a nocturnal decalcification of the reef community in summer.

Present-day conditions already imply reduced resilience to episodic

extreme events such as cyclones (Dove et al., 2013; see also Box

CC-CR).

448

Chapter 6 Ocean Systems

6.3.5.4. Ecosystems

The cumulative impacts of climate change drivers underlie alterations of

species interactions and ecosystem structure and functioning, including

changes in trophodynamics and the physical and chemical characteristics

of habitats (high confidence). These effects combine with more indirect

effects, such as shifts in stratification and productivity, expanding oxygen

minimum zones, and the changing composition and biomass of food

(partly resulting from direct effects on prey organisms) (high confidence).

These complexities reduce the precision and reliability of quantitative

projections (Section 6.5), including uncertainties concerning shifts in

upwelling and their future role in global primary production and the

development of fish stocks (Box CC-UP).

At the level of animal communities, effects of various drivers remain

largely unexplored, some are highly complex. For example, the net

eastward shift of Pacific skipjack tuna between 1980 and 2009 was linked

to the shifting aggregation of macrozooplankton and micronekton,

involving complex interactions of climate variability (due to ENSO;

Section 30.5.2), warming ocean surface, shallowing mixed layer depth

relative to the position of the warm pool, and the convergence of the

pool with the Pacific Equatorial Divergence Province (Lehodey et al.,

2011; see also Section 30.5.6.1.1). Interactive drivers will affect the

relative performance of interacting species, thereby shifting species

ranges, interactions, and food webs (medium confidence; Figure 6-7a).

Adaptation to various climate zones modifies the roles of light and

temperature in seasonalities and species interactions (Bradshaw and

Holzapfel, 2010). Moderate hypoxia expansion in warming seas, for

example, as the stratified central North Sea (Queste et al., 2013) may

well influence the degree of temperature-induced species displacements

(Figure 6-7b).

Impacts of climate change on benthic ecosystem engineers can also

profoundly alter ecosystems. Tropical corals respond to ocean warming

and acidification by increased bleaching, impeded calcification rates,

and increased incidence of disease (high confidence; Veron et al., 2009;

Veron, 2011; see also Sections 6.3.1-2, 30.5.6; Box CC-CR). In coral reefs

under multiple stressors, differentiation of these large-scale phenomena

into species-specific sensitivities is highly uncertain as trend data are

virtually nonexistent (Brainard et al., 2011). Little is known about impacts

on deep-water or cold-water corals and sponges, tropical calcified algae,

bryozoans, sponges, and tube-forming serpulid worms (Wood, 1999).

The reliance of all of these on surface productivity makes them vulnerable

to any alteration in food supply. Projected severe stress from increased

temperature, hypoxia, and ocean acidification will cause reduced

performance and increasing mortality in ecosystem engineers (high

confidence), and a deterioration of habitat characteristics for other

organisms (medium to low confidence).

As a corollary, shifts in the geographical distributions of marine species

(e.g., to higher latitudes or deeper waters; Figure 6-7b; Section 6.5.2)

cause changes in community composition and interactions (Harley,

2011; Simpson et al., 2011; Hazen et al., 2013). Some species may gain

predominance and abundance from fitness benefits (Figure 6-7) while

others become less competitive or easier prey (Occhipinti-Ambrogi,

2007). Thereby, climate change will reassemble communities and affect

biodiversity, with differences over time and between biomes and

atitudes (high confidence; Parmesan and Matthews, 2005; Sala and

Knowlton, 2006; Cheung et al., 2009; Parmesan et al., 2011; see also

Box CC-PP; Section 6.5).

6.3.6. Food Web Consequences

Community reassembly under climate change involves a change in

species composition and strongly alters food web structure, for example,

causing shifts in trophic pathways (Kirby and Beaugrand, 2009; Moloney

et al., 2011; see also Figure 6-12), some of which are irreversible (Jarre

and Shannon, 2010). Through trophic cascades (Cury et al., 2003; Luczak

et al., 2011), climate affects predation, competition, and food availability

(e.g., via changes in NPP; Figure 6-12; Utne-Palm et al., 2010), including

fish stocks (Parsons and Lear, 2001; Brown et al., 2010). Trophic

amplification then drives an ecosystem towards a new stable structure

or regime, which may be difficult to reverse (Folke et al., 2004). Warming

may result in consumer control of food web structure because respiration

of heterotrophic zooplankton and bacteria increases more strongly with

warming than does photosynthesis of autotrophic phytoplankton

(medium confidence; O’Connor et al., 2009).

Many impacts of climate change on food webs resemble those caused

by fishing, pollution, eutrophication, and associated hypoxia (Section

6.3.3), and habitat change (Brander, 2007); unambiguous attribution to

climate remains difficult (low to medium confidence; Parmesan et al.,

2011). Some of these factors also affect food web responses to climate

change. Fishing truncates the age and size structure of populations,

making them more dependent on annual recruitment and reducing their

ability to buffer environmental fluctuations (Genner et al., 2010; Planque

et al., 2010; Botsford et al., 2011; see also Figure 6-12). Both adult and

larval fishes show greater variability in abundance in exploited

compared to unexploited populations (Hsieh et al., 2008). Warming,

acidification, and removal of top or competing predators may all

contribute to large fluctuations in gelatinous plankton (e.g., jellyfish)

populations (low confidence; Molinero et al., 2005; Richardson and

Gibbons, 2008; Richardson et al., 2009; Condon et al., 2012).

Analyzing impacts on key species provides insight into how individual

components of a food web will respond to perturbations. However,

projections of future states must include the complex food web interactions

that influence the species and system-level responses, which affect

stability and resilience of the overall ecosystem (Neutel et al., 2007;

Dunne and Williams, 2009; Romanuk et al., 2009). There is no single

approach currently available that includes the complex links within and

among ecosystems, biogeochemistry, and climate as needed for projections

of future states of marine food webs (Fulton, 2011; Moloney et al., 2011).

In conclusion, there is low confidence in the quantitative projections of

such changes (for further discussion see Section 6.5).

6.3.7. Marine Reptiles, Mammals, and Birds

6.3.7.1. Principles

Marine reptiles (turtles, snakes, crocodiles), mammals, and seabirds

breathe air but live mostly in water; some shift or expand their ranges

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Ocean Systems Chapter 6

as a result of climate warming. The body temperature of ectothermic

reptiles is set by ambient conditions; only at large body size may their

body store heat and its temperature be higher than ambient. Reptiles

are thus more responsive to temperature than homeothermic seabirds

and marine mammals (McMahon and Hays, 2006), which regulate their

body temperature by adjusting metabolic heat production and insulation

from the environment, a trait beneficial especially in the cold. Various

degrees of body core insulation in mammals and birds constrain their

distribution to either warmer or colder waters (by poor or high insulation,

respectively). However, large body sizes enable some aquatic air

breathers to travel across the widest temperature ranges possible in

some of the largest migrations on Earth.

Changes in water chemistry and hypoxia have minimal direct influences

on the air-breathing vertebrates, reflecting their large independence

from physical and chemical drivers in the oceans. There is evidence for

increased sound propagation in a CO

-enriched ocean, but no evidence

yet for any effect on biota (Ilyina et al., 2010). If habitat structures

offering retreat or ambush disappear, this will increase the energetic

costs of life. Warming waters increase the cost of pursuit-diving as prey

fishes increase swimming velocity. The predation success of such

mammals (e.g., sea lions) and seabirds (e.g., penguins, cormorants) is

thus constrained to waters ≤20ºC (Cairns et al., 2008), a trend that

extrapolates into the future (low to medium confidence). As prey

distributions shift, foragers tied to land between trips may be

constrained by the physiological costs of finding prey (Péron et al., 2012;

Hazen et al., 2013). If food items are only found in thermally restricted

areas or move to greater depths, mammals and birds may become

constrained to certain distribution ranges or to the physiological limits

of their diving ability (McIntyre et al., 2011). Conversely, hypoxic habitat

compression for fishes may facilitate foraging opportunities for their

air-breathing predators (Hazen et al., 2009). Accordingly, many air-

breathers encounter changing habitat and food availability with climate

change (high confidence).

6.3.7.2. Field Observations

Some species of seabirds, marine mammals, and sea turtles have

responded to the anomalous ocean climate of the 20th century (high

confidence; Hughes, 2000). There is insufficient information to assess

effects on sea snakes or crocodiles. Poleward distribution shifts of turtles

consistent with recent warming have been recorded in almost all marine

groups. Decadal-scale climate fluctuations affect their recruitment

–1

–2

–3

–4

–5

–6

–7

–8

rimary production

Megafauna

Body Mass (g)

Small ﬁshes (sp. A)

Large ﬁshes (sp. A)

Large ﬁshes (sp. B)

Large ﬁshes (sp. C)

mall ﬁshes (sp. B)

Zooplankton and other invertebrates

Detritus / particulate

rganic matter

iogeographic shifts

ome species experience distribution shifts, abundance losses, or

become locally extinct (dashed lines). Loss and invasion of prey

organisms and/or predators may lead to changing diversity and

oodweb structure and result in trophic cascade effects.

henological shift

hanges in phenology due to climate change

ay lead to mismatch phenomena between

redator and prey, which may reduce

bundance of the predator.

hanges in body size

arine water-breathing ectotherms increase

heir consumption rate and have a smaller

aximum body size during warming. Changes in

he body size spectrum and food consumption

rate may lead to changes in foodweb structure

nd dynamics.

Large ﬁshes

(sp

(

. C

)

Predator–prey interactions Interactions that will be weakened or removed by climate changeCompetition

Figure 6-12 | Schematic diagram of expected responses to climate change in a marine food web. A coupled pelagic and benthic food web is structured by the body size

spectrum of species. Combined warming, hypoxia, and ocean acidiﬁcation reduce body size, shift biogeographies, change species composition and abundance, and reconﬁgure

trophic linkages and interaction dynamics. Fishing generally removes large-bodied species and truncates the body-size spectrum of the community. This confounds the detection

and attribution of food web responses to climate change. Arrows represent species interactions (e.g., between predator and prey or competitors for food or space). Broken lines

reﬂect the potential loss of populations and trophic linkages due to climate change.

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Chapter 6 Ocean Systems

uccess and nesting abundance (Van Houtan and Halley, 2011), with an

inverse correlation between warming and abundance in various species

and regions (Balazs and Chaloupka, 2004; Chaloupka et al., 2008;

Mazaris et al., 2009). Extreme weather causes nest flooding, considerably

reducing hatching success (Van Houtan and Bass, 2007); projected sea

level rise (WGI AR5 Chapter 13) will exacerbate such impact. Those with

high fidelity to nesting and foraging sites (Cuevas et al., 2008) are

impacted more than those capable of changing those sites (Fish et al.,

2009; Hawkes et al., 2009). Continued warming, modulated by changing

rainfall (Santidrián Tomillo et al., 2012), may skew turtle sex ratios

toward females, increase egg and hatchling mortality (Fuentes et al.,

2009), cause earlier onset of nesting (Pike et al., 2006; Mazaris et al.,

2008), decrease nesting populations (Chaloupka et al., 2008), and shift

dietary breadths (Hawkes et al., 2009), leading to projected recruitment

declines (e.g., leatherback turtles; Saba et al., 2012). Vulnerability due

to shifting sex ratio alone remains unclear, as nesting beaches have

persisted with low production of male hatchlings over decades or longer

(low confidence; Godfrey et al., 1999; Broderick et al., 2000; Hays et al.,

2003). The absence of sea turtles in certain regions may be best

explained by the temporal unavailability of food resources or strong

thermoclines restricting their bottom foraging abilities (Braun-McNeill

et al., 2008; Gardner et al., 2008).

Seabird range modifications probably caused by climate change were

recorded in polar areas and the temperate zone of the North Atlantic

(Grémillet and Boulinier, 2009). Temperate species have shifted their

ranges to higher latitudes in both hemispheres (Bunce et al., 2002;

Robinson et al., 2005; La Sorte and Jetz, 2010). Some species, like the

king penguin, follow shifting foraging zones (Péron et al., 2012); others,

such as the emperor penguin, are affected by changing habitat structure

(sea ice; Jenouvrier et al., 2012). Warming causes many bird species to

breed earlier (Sydeman and Bograd, 2009). High-latitude, cool-water

species undergo extended breeding seasons (Chambers et al., 2011). There

is often no agreement, whether changes reflect solely ocean warming, or

a combination of factors, such as fishing pressure on seabirds’ prey

species, sea level rise, and pollution (Galbraith et al., 2005; Votier et al.,

2005; Heath et al., 2009). Most shifts in range and seasonal activity

involve shifts in trophic relationships (medium confidence). Seabirds with

narrow geographic domains are expected to be more susceptible to

climate change (Chambers et al., 2005; Grémillet and Boulinier, 2009),

even leading to local extinctions (e.g., the Galápagos penguin: Vargas

et al., 2007; or the marbled murrelet: Becker et al., 2007).

The distribution, phenology, and migratory timing of marine mammals

are also shaped by predator-prey dynamics and climate impacts on

specific habitats (Calambokidis et al., 2009; Salvadeo et al., 2011). Some

marine mammals, that is, dolphin, porpoise, and whale species, shift their

distribution poleward to follow the movement of their prey (medium

confidence; Springer et al., 1999; MacLeod et al., 2005; Simmonds and

Isaac, 2007; Salvadeo et al., 2010 ). As in birds, vulnerability to climate

change is high for marine mammals with narrow geographic ranges

and high habitat dependence. For example, the critically endangered

vaquita, endemic to the Northern Gulf of California, cannot move north

because of the land barrier (MacLeod, 2009). The polar bear (Laidre et

al., 2008; Rode et al., 2012) and the walrus depend on sea ice as a

platform for hunting, resting, and giving birth. For polar bears, access

to prey such as ringed seals has been disrupted by the later formation

nd earlier breakup of sea ice in the eastern Canadian Arctic. Seasonal

migrants into the Arctic (fin, minke, gray, killer, humpback whales) may

increasingly compete with species adapted to operate in habitat with

sea ice (some seals, narwhal, bowhead whale, beluga). Both may benefit

from the net loss of sea ice, which will offer them better access to

foraging in a pelagic-dominated ecosystem (Moore and Huntington,

2008).

6.3.8. Summary and Conclusions

An organism’s capacity to perform, but also its access to food energy

fueling that performance, shape its sensitivity to climate change

(high confidence). Extreme temperatures surpassing the fringes of the

thermal envelope cause local abundance losses, extinction, and shifts

in temperature-dependent distribution ranges (high confidence; Section

6.3.1).

Some climate change effects detected in the field can be attributed to

temperature, but few allow clear attribution to other drivers (Sections

6.3.1-5, 6.6). In fishes and invertebrates, specialization in regional climate

regimes co-defines sensitivity to warming, acidification, and hypoxia

(high confidence; Section 6.3.5). In marine mammals, birds, and

ectothermic reptiles, changes in life history and population dynamics have

often not been directly attributed to climate drivers (low confidence), but

rather to the availability of habitat and food (high confidence; Section

6.3.7).

Natural climatic variability (Figure 6-1) and anthropogenic change, with

a strong role of warming, cause large-scale changes in biogeography,

abundance, diversity, community composition, and structure of marine

species (very high confidence; Section 6.3.1). Warming reduces body size

(medium confidence; Section 6.3.1). Differential species responses modify

their interactions across trophic levels through trophic amplification

(medium to high confidence; Section 6.3.6).

Some tropical species and ecosystems exist close to upper thermal limits

placing them among the marine ecosystems most affected by climate

change (high confidence; Section 6.3.1). Corals and coral reefs are

primary examples. However, other factors change concomitantly, such

that quantifying ecosystem changes attributable to warming or other

drivers has not always been possible (Section 6.3.5).

Under future climate change ocean acidification will affect marine

organisms and ecosystems for centuries (high confidence; Sections 6.3.2,

6.3.5). To date, very few ecosystem-level changes in the field have

been attributed to anthropogenic or local ocean acidification (medium

confidence; Section 6.3.2). Concomitant trends of warming, O

depletion,

OA, and other drivers prevent clear attribution to OA (Section 6.3.5).

Elevated CO

levels stimulate primary production of some macroalgae

and seagrass species (high confidence), causing them to be more

competitive than calcifying organisms (medium confidence; Section

6.3.2). High sensitivities to OA are associated with low capacities to

maintain pH in internal fluids (high confidence). Calcification rates in

sensitive invertebrates, including corals, echinoderms, and mollusks,

decrease under OA, especially if combined with temperature extremes

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Ocean Systems Chapter 6

(high confidence; Section 6.3.5). Thresholds beyond which effects occur

can be quantified only with low confidence; there are differential

sensitivities and thresholds between taxa and species (high confidence;

Section 6.3.2).

Expansion of oxygen minimum zones leads to community shifts clearly

attributable to extreme hypoxia (high confidence; Section 6.3.3). Gradual

effects of a progressive decline in ocean O

levels on communities have

not been sufficiently explored.

In general, community reassembly with new species coming in will occur

in the transition to future climates (medium confidence) and lead to

new ecosystem states (low confidence; Section 6.3.6). Climate change

interacts with top-down human interferences, such as fisheries or other

forms of harvesting, which accelerate impacts (medium confidence).

Nonlinearities challenge the projection of marine ecosystem trajectories

(FAQ 6.4).

In microbes, a conceptual foundation suitable to support an integrated

understanding of climate impacts on individual species and communities

is lacking. Specific physiological responses, such as in primary production,

fixation, or calcification, can be attributed to multiple environmental

drivers associated with climate change (high confidence; Sections 6.3.1-5).

6.4. Human Activities in Marine Ecosystems:

Adaptation Benefits and Threats

Human societies benefit from resources and processes supplied by marine

ecosystems, so-called ecosystem services. Attributing and projecting

Frequently Asked Questions

FAQ 6.4 | What changes in marine ecosystems are likely because of climate change?

There is general consensus among scientists that climate change signiﬁcantly affects marine ecosystems and may

have profound impacts on future ocean biodiversity. Recent changes in the distribution of species as well as species

richness within some marine communities and the structure of those communities have been attributed to ocean

warming. Projected changes in physical and biogeochemical drivers such as temperature, CO

content and acidiﬁcation,

oxygen levels, the availability of nutrients, and the amount of ocean covered by ice will affect marine life.

Overall, climate change will lead to large-scale shifts in the patterns of marine productivity, biodiversity, community

composition, and ecosystem structure. Regional extinction of species that are sensitive to climate change will lead

to a decrease in species richness. In particular, the impacts of climate change on vulnerable organisms such as warm-

water corals are expected to affect associated ecosystems, such as coral reef communities.

Ocean primary production of the phytoplankton at the base of the marine food chain is expected to change but

the global patterns of these changes are difﬁcult to project. Existing projections suggest an increase in primary

production at high latitudes such as the Arctic and Southern Oceans (because the amount of sunlight available for

photosynthesis of phytoplankton goes up as the amount of water covered by ice decreases). Decreases are projected

for ocean primary production in the tropics and at mid-latitudes because of reduced nutrient supply. Alteration of

the biology, distribution, and seasonal activity of marine organisms will disturb food web interactions such as the

grazing of copepods (tiny crustaceans) on planktonic algae, another important foundational level of the marine

food chain. Increasing temperature, nutrient ﬂuctuations, and human-induced eutrophication may support the

development of harmful algal blooms in coastal areas. Similar effects are expected in upwelling areas where wind

and currents bring colder and nutrient-rich water to the surface. Climate change may also cause shifts in the

distribution and abundance of pathogens such as those that cause cholera.

Most climate change scenarios foresee a shift or expansion of the ranges of many species of plankton, ﬁsh, and

invertebrates toward higher latitudes, by tens of kilometers per decade, contributing to changes in species richness

and altered community composition. Organisms less likely to shift to higher latitudes because they are more tolerant

of the direct effects of climate change or less mobile may also be affected because climate change will alter the

existing food webs on which they depend.

In polar areas, populations of species of invertebrates and ﬁsh adapted to colder waters may decline as they have

no place to go. Some of those species may face local extinction. Some species in semi-enclosed seas such as the

Wadden Sea and the Mediterranean Sea also face higher risk of local extinction because land boundaries around

those bodies of water will make it difﬁcult for those species to move laterally to escape waters that may be too

warm.

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Chapter 6 Ocean Systems

cosystem changes and their effects on human communities caused by

climate change including ocean acidification is challenging. Insufficient

observations compound an understanding of long-term changes and

the definition of baseline conditions. Some of the challenges are related

to the difficulty of projecting how human communities will adapt to

changing marine ecosystem benefits.

6.4.1. Ecosystem Services

Marine ecosystem services (e.g., Chapter 5) include products (food,

fuel, biochemical resources), climate regulation and biogeochemical

processes (CO

uptake, carbon storage, microbial water purification),

coastal protection, provision of space and waterways for maritime

transport, cultural services (recreational and spiritual opportunities,

aesthetic enjoyment), and functions supporting all other ecosystem

services (nutrient cycling, photosynthesis, habitat creation). Most

components of the marine environment contribute to more than one

major category of ecosystem service: for example, ocean primary

productivity is classified as a supporting service, but it affects provisioning

services via changes in fisheries, generation of fossil fuel resources,

regulating services via the global carbon cycle and climate regulation,

and cultural services via the enjoyment of a healthy ecosystem. Rarely

has economic damage of climate change to a whole ecosystem been

evaluated and projected. The projected loss of tropical reef cover due to

ocean acidification under SRES A1 and B2 scenarios will cause damages

of US$870 and 528 billion (year 2000 value) by 2100, respectively (cost

rising with parallel economic growth; Brander et al., 2012; see also Box

CC-OA). Such loss is felt most strongly in the respective regions.

6.4.1.1. Food from the Sea

Fisheries provide 3 billion people with almost 20% of their average per

capita intake of animal protein (FAO, 2012a), 400 million depend critically

on fish for their food (Garcia and Rosenberg, 2010). Total world marine

capture fisheries catches stabilized in the mid-1990s at about 90

milliontons per year. Marine aquaculture of primarily mollusks and

crustaceans contributes more than 63 million tons annually to seafood

production, mostly concentrated in coastal areas (FAO, 2012b). The growth

of aquaculture has decelerated, but is still considered a development

opportunity and a strong need in regions such as Africa and Latin America

(Section 7.4.2.2).

Climate-induced shifts in ecosystems and fisheries production will

create significant challenges to sustainability and management (Section

7.5.1.1.3), particularly for countries with fewer resources and lower

adaptive capacity, including many low-latitude and small island nations

(high confidence; Allison et al., 2009; Worm et al., 2009; Cooley et al.,

2012; see also Sections 7.2.1.2, 7.4.2.1, 30.6.2; WGIII AR5 Section 2.1).

Vulnerability will be exacerbated by increases in the frequency and

severity of extreme events (e.g., floods or storms) damaging infrastructure,

homes, health, livelihoods, or non-marine food security (Kovats et al.,

2003; Rosegrant and Cline, 2003; Adger et al., 2005; Haines et al., 2006).

The projected trends in fish stocks will widen the disparity in food

security between developing and developed nations. Fish migrations

ue to warming (Section 6.3.1) have already shifted the composition

of fisheries catches (Pinsky and Fogarty, 2012; Cheung et al., 2013a)

and altered stock distributions (Sabatés et al., 2006). Further warming

may be beneficial for fisheries productivity in some regions such as the

North Atlantic, because of the poleward shift of exploited species and

changes in primary productivity (Arnason, 2007; Stenevik and Sundby,

2007; Cheung et al., 2010; see also Box 6-1; Section 30.5.1.1.1), or for

some Pacific Islands due to the eastward redistribution of tuna stocks

(Lehodey, 2000; Lehodey et al., 2011). Resulting changes in accessibility

and fishing operations costs are projected to straddle economic zones,

perturb international fishery agreements, and cause excessive exploitation

(Hannesson, 2007; Sumaila et al., 2011; see also Sections 7.3.2.4, 7.4.2;

WGIII AR5 Section 4.3.7).

Invertebrate fisheries and aquaculture appear very vulnerable to the

impacts of ocean acidification (Barton et al., 2012; see also Box CC-OA;

Figure 6-10). This concerns especially shelled mollusks, with a substantial

decline in their global production projected between 2020 and 2060

under the SRES A2 business-as-usual scenario (Cooley and Doney, 2009;

Cooley et al., 2012). Effects on calcifying plankton will propagate

through the food web, making estimates of economic impact on fish catch

by OA difficult, also due to complex interactions with other stressors

like warming and fisheries management (Griffith et al., 2012; Branch et

al., 2013). Model projections suggest a potential loss of up to 13%

(SRES A1FI scenario) to annual total fishery value in the USA, or globally

more than US$100 billion annually by 2100 (Cooley and Doney, 2009;

Narita et al., 2012). Vulnerability differs highly between nations according

to the contribution of such fisheries to their economy (Cooley et al.,

2012; see also Sections 7.3.2.4, 7.4.2). These projections are sensitive

to the projected vulnerabilities of the organisms to ocean acidification

(medium confidence; Section 6.3.2).

Fishing reduces abundances at high trophic levels, but increases

abundances at mid-trophic levels. It reduces species numbers, simplifies

ecosystem structure, and increases ecosystem sensitivity to climate

change (Perry et al., 2010). Exploitation of fish stocks and the alteration of

their demography, population dynamics, and life history traits (Petitgas

et al., 2006; Perry et al., 2010; Planque et al., 2010) can reduce the

capacity of fish populations to buffer changes in climate variability

(Ottersen et al., 2006; Genner et al., 2010), and increase variability in

population size. Interactions between warming, OA, and human activities

such as fishing may thus exacerbate climate impacts on a wide range

of ocean processes and services, including marine fisheries (medium

confidence; Tables 6-4, 6-6; Section 30.6.2).

A 2°C global temperature increase by 2050 is estimated to cause

globallosses in landed value of US$17 to 41 billion annually (in 2005

value), with an estimated cost of adaptation for the fisheries of US$7

to 30 billion annually over a 40-year time frame between 2010 and

2050. The largest loss in landed value is projected to occur in East Asia

and the Pacific (low confidence;Sumaila and Cheung,2010). Overall

impacts and the regional manifestations will partially depend on the

flexibility and response capacities of food production systems (Elmqvist

et al., 2003; Planque et al., 2011a).

Specific implications for the fishing industry are still poorly known, as

future projections of shifts in primary production and knock-on effects

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Ocean Systems Chapter 6

hrough food webs and into fisheries remain uncertain (low confidence

in effects of changing NPP; Planque et al., 2011b; Stock et al., 2011).

6.4.1.2. Other Provisioning Services

Reductions in marine biodiversity due to climate change and other

anthropogenic stressors (Tittensor et al., 2010), such as OA (CBD, 2009)

and pollution, might reduce the discovery of genetic resources from

marine species useful in pharmaceutical, aquaculture, agriculture, and

other industries (Arrieta et al., 2010), leading to a loss of option value

from marine ecosystems. Climate change increases the demand for

marine renewable energy such as wind and wave power, though with

potential ecosystem impacts of their infrastructure (Section 6.4.2).

6.4.1.3. Climate Regulation and Extreme Events

The effect of climate change on marine biota will alter their contribution

to climate regulation, that is, the maintenance of the chemical composition

and physical processes in the atmosphere and oceans (high confidence;

Beaumont et al., 2007). Regulatory mechanisms in which organisms

(especially phytoplankton) play a key role, include control of the level

of atmospheric CO

through the balance between photosynthesis and

respiration (Johnson et al., 2010), and through the biological and alkalinity

pump (Falkowski, 1997; Feely et al., 2008). They also include the

modulation of further greenhouse gases such as nitrous oxide (N

Jin and Gruber, 2003; Law, 2008; see also Section 6.1.1.3), and the

modulation of other climatically reactive gases such as dimethylsulfide

(DMS; Vogt et al., 2008). A projected decrease in global ocean NPP

(Section 6.5.1) may result in decreased export of biogenic carbon to the

deep ocean (Bopp et al., 2002; Boyd and Doney, 2002; Hashioka and

Yamanaka, 2007). A positive feedback on climate change may result;

however, many of the factors controlling the pump are poorly understood

(Figure 6-4; WGI AR5 Chapter 6).

Coastal marine ecosystems reduce the effects of floods and storm

surges which account for most of the natural disasters affecting people

in coastal regions (IPCC, 2012a). Empirical and modeling studies show

that coral reefs contribute to buffering the impact of tsunamis (Fernando

et al., 2005; Gravelle and Mimura, 2008; see also Sections 5.4.2.4, 30.5;

Box CC-CR). Experiments and models indicate that warming and OA

slow coral growth by nearly 50% by 2050 (Box CC-CR; Section 5.4.2.4),

making some islands and coastal areas more vulnerable to tsunamis,

storm surges, wave energy, and coastal erosion (high confidence).

Wetlands and mangroves provide biologically diverse buffer zones

(Section 5.4.2.3). The combined impacts of climate change, pollution,

deoxygenation, and other overlapping stressors, on mangroves and

wetlands have not been determined (Cooley et al., 2009; Cooley, 2012).

Some of these stressors enhance each other’s effects in coastal systems

(Feely et al., 2010; Cai et al., 2011; Howarth et al., 2011).

6.4.1.4. Cultural Services

Cultural services encompass a wide array of services with marine

biodiversity as a core component supporting recreation and tourism as

he economically most relevant. Tropical coral reefs and their enormous

biodiversity sustain substantial tourist industries, presently with global

annual net benefits of about US$9.6 billion (Cesar et al., 2003; see also

Box CC-CR; Section 30.6.2.2). If reef services degrade, coastal visitors

might choose alternative attractions (UNWTO, 2008). Increased travel

to see disappearing ecosystem types (e.g., Antarctica: Liggett et al.,

2011) or in previously inhospitable areas or seasons (Amelung et al.,

2007; Moore, 2010) create new pressures and are unsustainable as the

locations of key attractors shift (e.g., cetaceans: Lambert et al., 2010;

Salvadeo et al., 2013).

Climate change may endanger harvests of marine species with spiritual

and aesthetic importance to indigenous cultures, raising ethical

questions about cultural preservation (e.g., Nuttall, 1998). In coastal

communities, losing the aesthetic values of marine ecosystems may

harm local economies: better water quality and fewer harmful algal

blooms are related to higher shellfish landings and real estate prices

(Jin et al., 2008).

Some heritage benefits of preserving marine ecosystems consist of the

economic value of a healthy, diverse ecosystem to future generations.

Any climate-related biodiversity loss or pollution of marine ecosystems

would decrease the bank of resources for future opportunities. For

example, the research and conservation value of coral reef biodiversity

and its non-use value are estimated together at US$5.5 billion annually

(Cesar et al., 2003). As with spiritual and aesthetic benefits, maintaining

heritage benefits under climate change poses challenges for managers

concerning equity and ethics as well as multigenerational (and possibly

multi-cultural) ethical questions.

6.4.1.5. Supporting Services

Fully identifying the services supporting other ecosystem benefits is

virtually impossible, as they are diverse in nature and scale. Ecosystem

engineers play an important role in these services. Damage to calcifying

algae and corals will reduce habitat for other species (Section 6.3.5),

biodiversity, cultural and leisure values, and their climate regulation

capacity.

Waterways for shipping are expected to change in the next several

decades (very high confidence; Chapter 28; Section 30.6.2.3). Reductions

in Arctic sea ice allow new trade routes such as the Northwest Passage

(Wilson et al., 2004; Granier et al., 2006), enabling economically viable

trans-Arctic shipping, and access to regional resources for exploitation

and tourism. This development would increase emission of greenhouse

gases and other pollutants (Lauer et al., 2009; Corbett et al., 2010), and

facilitate the invasion of non-indigenous species carried on hulls and in

ballast waters (Lewis et al., 2004).

6.4.2. Management-Related Adaptations and Risks

6.4.2.1. Ecosystem Management

A changing climate will have both positive and negative consequences

for managing ocean resources (high confidence) (Eide and Heen, 2002;

454

Chapter 6 Ocean Systems

ide, 2007; see also Section 6.4.1). Ecosystem-based management

(EBM, an approach recognizing all, including human interactions,

within an ecosystem) or the ecosystem approach (EA, a strategy for the

integrated management of living resources promoting both conservation

and sustainable use) are increasingly adopted globally (FAO, 2003) to

deal with the multitude of human pressures on marine ecosystems

(Sherman et al., 2005; Hoel, 2009). Extended EBM addresses changes

driven by climate and human activities, considering that diverse drivers

will interact and confound each other (Planque et al., 2010; Eero et al.,

2011; see also Section 6.3.5). Human activities will undermine resilience

to other, including climate, impacts or undermine the effectiveness of

mitigation and adaptation measures, by increasing variability (thereby

reducing predictability), and limiting scope for adaptation (high confidence;

e.g., Hughes, 2004; Sissener and Bjørndal, 2005; Eero et al., 2011). Thus,

managing ecosystems under climate change increases the resilience of

ecosystems and adaptive capacity of management systems through

reducing other human perturbations (e.g., overfishing) (Brander, 2008;

see also Section 7.5.1.1.3). Managing ecosystems also reduces the

consequences of ocean acidification until CO

emission reduction becomes

effective (Rau et al., 2012; Billé et al., 2013; McLeod et al., 2013; see

also Box CC-OA). Ecosystem resilience is enhanced by reducing regional

eutrophication (Falkenberg et al., 2013), or in aquaculture by avoiding

acidified water (Barton et al., 2012) and by selecting and cultivating

pre-adapted strains (Parker et al., 2012).

However, effects of climate change cannot be reversed by reducing the

impacts of non-climatic drivers, emphasizing the need for adaptive

management. Increased variability of ecosystem responses to climate

change and the low predictability of some biological responses

undermine the effectiveness of management and conservation

measures. A particular risk is that climate change may contribute to

large-scale ecosystem regime shifts (Section 6.3.1.5; Box 6-1). Detecting

and forecasting such shifts from time series of environmental and

biological data (Carpenter and Brock, 2006; deYoung et al., 2008), is

constrained by an insufficient number of observations and limited

quantitative understanding (Section 6.1.2). Biogeographic shifts

challenge spatial management (Box CC-MB; Sections 6.3.1, 6.5), which

is a fundamental part of EBM (Douvere, 2008), and demand that “fixed

in law forever” site-attached zoning to protect specific species may

need to become more flexible to maintain the original objectives as

species move or community structures shift (high confidence; Soto,

2001; Hawkins, 2012).

6.4.2.2. Geoengineering Approaches

Geoengineering approaches to mitigate climate change and its effects,

include Solar Radiation Management (SRM) and Carbon Dioxide

Removal (CDR; see Table 6-5; IPCC, 2012b). SRM aims to reduce warming

by increasing albedo, for example, via stratospheric injection of sulfate

aerosol (Crutzen, 2006). SRM may affect marine ecosystems through

changes in precipitation. With continued CO

emissions it leaves ocean

acidification largely unabated as it cannot mitigate rising atmospheric

concentrations (Vaughan and Lenton, 2011; Williamson and Turley,

2012). Termination of SRM after its implementation involves the risk of

rapid climate change and more severe effects on ecosystems (Russell

et al., 2012).

roposed CDR techniques include both ocean- and land-based

approaches (Vaughan and Lenton, 2011; see also Section 30.6.4). CO

storage in geological reservoirs may occur beneath the seafloor, for

example, in porous marine aquifers, and includes the risk of CO

leakage

to the marine environment. Proposals to directly or indirectly sequester

into the ocean (Caldeira et al., 2005; Boyd, 2008; Shepherd et al.,

2009; see also Table 6-5; WGIII AR5 Section 7.5.5) include, among others,

the use of ocean fertilization techniques by nutrient addition, the direct

storage of biomass in the deep ocean, the addition of alkalinity for

build-up of dissolved inorganic carbon (DIC; i.e., carbonate), and the

direct CO

injection into the deep ocean (Williamson et al., 2012). All of

these approaches have potentially negative consequences for marine

ecosystems.

Ocean fertilization by adding iron to high-nutrient low-chlorophyll

(HNLC) oceanic waters could increase productivity and the net export of

organic material to the deep ocean and its consecutive decomposition,

causing deep-water accumulation of CO

. Fertilization would affect all

major marine biogeochemical cycles of the ocean with unclear side

effects that could include the formation of methane (CH

) and N

O (Law,

2008) or the stimulation of harmful algal blooms (Trick et al., 2010). The

enhanced NPP would add more carbon to the base of food webs (de

Baar et al., 2005) and stimulate growth, for example, of deep-sea benthos

(Wolff et al., 2011). Any regional increase in organic material (through

fertilization or intentional storage of biomass) would cause enhanced

demand and deep-water O

depletion (Sarmiento et al., 2010; Table

6-5), increasing the level and extent of hypoxia and associated impacts

on marine ecosystems (Sections 6.3.3, 6.3.5, 30.5.7). The synergistic

effects of CO

-induced acidification will exacerbate the biological

impacts (high confidence).

Neutralizing the acidifying water by the addition of alkalinity, for

example, calcium oxide, would require large-scale terrestrial mining

with associated consequences (Caldeira et al., 2005). The biological

effects of increased concentrations of Ca

ions and dissolved inorganic

carbon remain insufficiently explored. Direct injection of CO

or its

localized disposal in the ocean (e.g., as a lake in a deep-sea valley)

causes locally highly increased CO

and acidification effects on deep-

sea organisms (high confidence; Caldeira et al., 2005; see also Section

6.3.3.4). In contrast to long-term ocean fertilization or storage of

biomass, this technique leaves the oxygen inventory of the deep ocean

untouched (limited evidence, medium agreement; Pörtner et al., 2005).

The knowledge base on the implementation of SRM and CDR techniques

and associated risks is presently insufficient. Comparative assessments

suggest that the main ocean-related geoengineering approaches are

very costly and have large environmental footprints (high confidence;

Boyd, 2008; Vaughan and Lenton, 2011; Russell et al., 2012).

6.4.2.3. Health Issues

Human health and near-shore ecosystems may be directly impacted by

climate change effects on harmful algal blooms (HABs; Edwards et al.,

2006; see also Section 30.6.3) or disease vectors. Planktonic time-series

archives and nearshore sediment cores containing HAB cysts have

revealed few examples of strong linkages between altered HABs and

455

Ocean Systems Chapter 6

climate fluctuations (Dale et al., 2006; see also Section 30.5.3.1.2). HABs

can be stimulated by warming, nutrient fluctuations in upwelling areas,

eutrophication in coastal areas, and enhanced surface stratification

(medium confidence). Species-specific responses involve shifts in seasonal

cycles and blooms (Johns et al., 2003). Ocean acidification may exacerbate

the toxicity of species in coastal oceans under nutrient-limited conditions

(Tatters et al., 2012; Sun et al., 2011). Suitable adaptation measures

include appropriate monitoring of biotoxin problems (Hallegraeff, 2010).

Continued warming of tropical and temperate coastal habitats, excessive

nutrient loading leading to phytoplankton and zooplankton blooms,

and sea water inundation due to sea level rise are all projected to

exacerbate the expansion and threat of cholera (medium confidence;

see also Sections 11.5.2.1, 30.6.3), although attribution to climate

change is confounded by climate variability and non-climate drivers

(Lafferty, 2009; Dobson, 2009).

Cholera and its pathogen, the marine bacterium, Vibrio cholera, have

been widely studied. The pathogen associates with marine organisms,

especially chitinized zooplankton (Vezzulli et al., 2010). Where cholera

is endemic (e.g., India, Bangladesh, Latin America), outbreaks correlate

with warming and high zooplankton abundance (Lobitz et al., 2000;

Lipp et al., 2002). Based on an 18-year climate record for Bangladesh,

Pascual et al. (2000) reported cholera outbreaks at ENSO events, and

the recent reappearance of cholera in Peru has also been linked to the

intense 1991–1992 ENSO (Lipp et al., 2002). An increase in sustained

maximum temperatures of the Baltic Sea (Section 30.5.3.1.4) has been

related to an increase in reported Vibrio infections; highest human

mortality rates were associated with V. vulnificus infections (Baker-

Austin et al., 2013). Continued warming of tropical and temperate

coastal habitats, excessive nutrient loading leading to phytoplankton

and zooplankton blooms, and seawater inundation due to sea level rise

are all projected to exacerbate the expansion and threat of cholera

(medium confidence).

Ciguatera poisoning may occur when people consume fish, mainly from

tropical reefs, that have ciguatoxins from the epiphytic dinoflagellate

Gambierdiscus sp. Historical records show significant correlations

between ciguatera poisoning and sea surface temperature in South

Pacific nations (Hales et al., 1999). However, the relationship is nonlinear

and dependent on the thermal window of the specific dinoflagellate

(Llewellyn, 2010). This casts doubt on the accuracy of projected increases

in ciguatera poisoning using linear extrapolations from observations

(low confidence).

Topic Brief description Challenge and impact References

Solar radiation management

echniques

Deﬂ ection of approximately 1.8% of sunlight, by

arious techniques, is able to offset the global mean

emperature effects of a doubling of atmospheric CO

content from pre-industrial values.

Will leave ocean acidiﬁ cation unabated (high

onﬁ dence). Response of primary production to light

eduction unclear.

Crutzen (2006); Caldeira and Wood (2008)

cean storage by direct

injection

apture of CO

post-combustion from mainly coastal

power plants, followed by injection of liquid CO

ipeline or from a ship into the deep ocean.

ill add to ocean acidiﬁ cation and create localized

harm to marine life (high conﬁ dence). Quantities will

e small relative to the atmospheric invasion signal.

injected will dissolve and be transported by ocean

irculation with eventual surface exposure.

aldeira et al. (2005)

Sub-sea geologic storage Capture of CO

from extracted gas or from post-

combustion followed by well injection into a porous

ubmarine aquifer beneath impermeable geologic

strata.

Extensive experience in place from the Norwegian

Sleipner ﬁ eld activity in the North Sea. No evidence of

cean impact from leakage to date.

Benson et al. (2005)

cean fertilization Spreading of trace amounts of reduced iron over very

large areas of the surface ocean where excess nutrients

ccur. Overcoming the local iron deﬁ ciency creates

extensive phytoplankton blooms drawing down sea

surface pCO

. Fertilization can also be carried out

y using direct or indirect (ocean pipes) addition of

macronutrients to oceanic regions where they are

epleted.

uch of the exported organic matter is remineralized

at shallow depths, creating local oxygen stress and

hallow CO

enrichment and methane and N

production. These effects are temporary and the

effective retention time is short. If sustained, reduced

urface ocean and increased deep ocean acidiﬁ cation.

loss in ocean interior (medium conﬁ dence).

e Baar et al. (1995); de Baar et al. (2005);

Pörtner et al. (2005); Boyd et al. (2007);

uesseler et al. (2008); Law (2008); Cao

and Caldeira (2010)

Artiﬁ cial upwelling or

ownwelling

Ocean fertilization by bringing nutrient rich deep water

(

from 200 to 1000 m) to the surface. Downwelling

occurs in parallel, transporting physically dissolved CO

into the deep ocean.

Deep water contains high levels of CO

, which if

eleased counteracts the binding of CO

by fertilization.

No evidence available.

Lovelock and Rapley (2007); Oschlies et

l. (2010)

equestration of organic

carbon

torage of terrestrial biomass in the coastal or deep

ocean.

hysical impact, regional loss of oxygen, CO

accumulation and acidiﬁ cation during degradation;

increases in methane, N

O, and H

S. No evidence

available.

etzger and Benford (2001); Strand and

Benford (2009)

Carbonate neutralization Dissolution of power plant ﬂ ue gas into sea water

yielding an acidic solution that is neutralized

by addition of crushed limestone. The resulting

bicarbonate-rich ﬂ uid is discharged to the ocean.

Involves the transport and crushing to ﬁ ne scale of

large quantities of limestone and the processing of very

large quantities of sea water. Environmental impact

issues not yet explored.

Rau (2011)

Accelerated olivine

weathering

Uses wind powered electrochemical processes to

remove HCl from the ocean and neutralizes the acid

with silicate minerals such as olivine for disposal. The

net result is to add alkalinity to the ocean akin to

natural silicate weathering processes.

Complex system as yet untested in pilot processes.

Involves mining and crushing large quantities of silicate

minerals. Very long time scale consequences uncertain.

House et al. (2007); Köhler et al. (2010)

Table 6-5 | Challenges for the oceans that will arise from the employment of a range of geoengineering methods (SRM = solar radiation management; CDR = carbon dioxide

removal).

456

Chapter 6 Ocean Systems

6.4.3. Conclusions

Human societies benefit from and depend on marine ecosystem services,

including the provisioning of food and other goods, regulation of climate

and extreme events, and cultural and supporting services (Section

6.4.1). Attributing and projecting climate-change-mediated shifts in

these services remains a challenge, due to the intrinsic difficulty of

assessments, lack of baseline and long time series data, and confounding

human impacts. However, empirical and modeling studies indicate that

climate change impacts on marine ecosystems lead to changes in

provisioning, regulating, and supporting services (high confidence), as

well as cultural services (limited evidence, medium agreement).

Food production from the sea is facing diverse stressors (Section 6.4.1.1),

such as overfishing and habitat degradation, which interact with climate

change phenomena, including warming (Section 6.3.1), ocean acidification

(Section 6.3.2), and hypoxia (Section 6.3.3). Projections of impacts on

capture fisheries are constrained by uncertainties in marine primary

production (medium evidence, medium agreement; Section 6.5.1).

Negative effects are projected to be most significant in developing

nations in tropical regions (high confidence). Nations at higher latitudes

may even benefit from climate change effects on ocean ecosystems, at

least initially (Section 6.5.3).

Climate change effects on biota will alter their climate regulation

through mechanisms such as carbonate production, the biological

pump, the balance between photosynthesis and respiration, and the

modulation of greenhouse gases (high confidence; Section 6.4.1.3).

However, projections of the direction and magnitudes of feedbacks are

at an early stage (low confidence).

Future management of ecosystems and fisheries might have to aim for

increasing ecosystem resilience to climate change, for example, through

reductions of other human perturbations (Section 6.4.2.1). Active ocean

geoengineering strategies to ameliorate climate change may prove

detrimental to the functioning of ecosystems, which highlights the need

for further research and careful governance (Section 6.4.2.2). There is

limited understanding of how harmful algal blooms and pathogens

affecting human health will respond to climate change (Section 6.4.2.3;

medium to low confidence).

6.5. Projections of Future Climate Change

Impacts through Modeling Approaches

A range of models explore climate change effects on marine biota, from

primary producers to higher trophic levels, and test hypotheses about

responses of marine species, food webs, and ecosystems (Rose et al.,

2010; Fulton et al., 2011; Stock et al., 2011; see also FAQ 6.2). Both

empirical and mechanistic approaches are used over a range of temporal

and spatial scales (Barange et al., 2010; Stock et al., 2011). There is an

increasing need for upscaling from molecular and physiological to

ecosystem level (e.g., Le Quesne and Pinnegar, 2012). Uncertainty in

projections of changes in marine ecosystems is partly contingent on the

level of confidence in climatic and oceanographic projections (Section

6.1.1; WGI AR5 Section 9.8). Models are currently useful for developing

scenarios of directional changes in net primary productivity, species

istributions, community structure, and trophic dynamics of marine

ecosystems, as well as their implications for ecosystem goods and services

under climate change. However, specific quantitative projections by

these models remain imprecise (low confidence; Hannah et al., 2010;

Rose et al., 2010; Stock et al., 2011; FAQ 6.4).

Earth System Models couple atmosphere, cryosphere, and hydrosphere

(including the oceans), as well as climate and carbon cycles, and project

changes in ocean biogeochemistry under a range of CO

emission scenarios

(WGI AR5 Chapter 6). Models focusing on population and species level

responses comprise models of population dynamics, models of species

distribution, and models which explicitly link effects of changes in ocean

physics and chemistry to changes in interactions between species at

different trophic levels, or human activities such as fishing and

aquaculture (Rose et al., 2010).

6.5.1. Oceanic Primary Production

Climate-induced effects on global ocean NPP comprise changes in its

long-term average, seasonal timing, and peak amplitude (Henson et al.,

2013). The magnitude, direction, and pattern of projected changes vary

with differences in model structure and parameterization (Box CC-PP;

Figure 6-13). Unknown accuracy of current NPP observations further

increases the uncertainty of projections, as does the incomplete

understanding of effects of multiple drivers on NPP (Sections 6.3.1-5,

6.4). Global coupled climate-ocean biogeochemical Earth System

Models (WGI AR5 Chapter 6) project an increase in NPP at high latitudes

but a decrease in permanently stratified oceans at mid-latitudes, in the

tropics (west tropical Pacific, tropical Indian Ocean, tropical Atlantic),

and in the North Atlantic (medium confidence; Steinacher et al., 2010;

Bopp et al., 2013) (Figure 6-13). The overall result is a reduction in global

mean NPP under all RCP scenarios (medium confidence in the direction

of projected trends, low confidence in the magnitude of change).

6.5.2. Higher Trophic Levels

Projected future changes in temperature and other physical and

chemical oceanographic factors are expected to affect the distribution

and abundance of marine fishes and invertebrates, as elaborated by

species distribution models. Limits of distribution ranges of 1066 exploited

species are projected to undergo shifts by a median of around 50 km

per decade to higher latitudes by 2050 relative to 2000 under the SRES

A1B (≈RCP6.0) scenario (Cheung et al., 2009). Some species shift

toward the equator following a regional temperature gradient (Burrows

et al., 2011; Cheung et al., 2013b; Pinsky et al., 2013). The rate of range

shifts is projected to be three times higher for pelagic than for demersal

fishes (Cheung et al., 2009), the latter shifting at a rate of around 27 to

36 km per decade (Cheung et al., 2013b). However, the expansion of

hypoxic waters may have a greater impact than warming on demersal

fishes (Koslow et al., 2011). As a result of distribution shifts, high-

latitude regions (the Arctic, Southern Ocean) are projected to have high

rates of species invasions. Intermediate latitudes are expected to undergo

both invasions and local extinctions. High rates of local extinction are

projected for the tropics and semi-enclosed seas (e.g., Mediterranean

Sea, Persian Gulf). In addition, the future productivity and distribution

457

Ocean Systems Chapter 6

of higher trophic level organisms are projected to change due to changes

in primary productivity (Section 6.3.6). For example, the migration route

of Pacific sardine is projected to shift because of changes in primary

productivity and food availability (Ito et al., 2010). The global pattern

of distribution shifts is generally consistent with regional-scale projections

and past observations (e.g., Lenoir et al., 2011; Cheung et al., 2013a).

However, detailed quantitative projections are sensitive to model structure

and assumptions (Hare et al., 2012; Jones et al., 2013) and responses

of specific populations may differ from average species responses (Hazen

et al., 2013).

Coral reefs are projected to undergo long-term degradation by 2020 to

2100 relative to the 2000s under RCP2.6, 4.5, and 8.5 or their equivalents

(Section 30.5.6). Reefs projected to be threatened most by bleaching

under the SRES A1B scenario by 2100 include the Central and Western

Equatorial Pacific, Coral Triangle, and parts of Micronesia and Melanesia

(Teneva et al., 2012). These projections assume that coral bleaching

occurs when SST exceeds a certain threshold, and that there is limited

potential to shift such threshold by adaptation. Reef degradation will

impact ecosystem services (Hoegh-Guldberg, 2011; see also Section 6.4;

Box CC-CR).

Some groups of marine air-breathing fauna are projected to shift in

distribution and abundance (Section 6.3.7). Cetacean richness will

increase above 40° latitude in both hemispheres, while at lower latitudes

both pinniped and cetacean richness are projected to decrease by 2040–

2049 relative to 1990–1999 under the SRES A1B scenario (Kaschner et

al., 2011). Using SST as a predictor, the distribution of loggerhead turtles

is projected to expand poleward in the Atlantic Ocean and to gain habitat

in the Mediterranean Sea by 2070–2089 relative to 1970–1989 (Witt

et al., 2010). Leatherback turtle may decrease in abundance at a rate

of 7% per decade because of reduced hatching success with warming

following the SRES A2 scenario (Saba et al., 2012). Abundances of some

seabirds such as European breeding seabirds (Huntley et al., 2007),

Cassin’s auklet in the California Current Ecosystem, or emperor penguin

in Antarctica are projected to decline because of climate-induced changes

in oceanographic conditions, such as temperature and upwelling intensity

(Wolf et al., 2010; see also Box CC-UP), or summer sea ice conditions

(Jenouvrier et al., 2012). The diversity of megafaunal responses to climate

change will have cascading ecosystem impacts, and will affect ecosystem

services such as tourism (high confidence; Sections 6.3.7, 6.4.1).

6.5.3. Ecosystems and Fisheries

One of the most direct impacts of climate change on marine ecosystem

services is through fisheries (Sections 6.4.1, 7.2.1.2, 7.3.2.4, 7.4.2).

Projected climate impacts on fisheries are based on recruitment, growth,

mortality, abundance, and distribution of fish stocks as well as changes in

ocean NPP (Cheung et al., 2008), evaluated from chlorophyll concentration

and other variables such as sea surface temperature (Campbell et al.,

2002). Friedland et al. (2012) suggested thatchlorophyll concentration,

indicating both phytoplankton production and biomass, is a better

predictor of the fishery yield in large marine ecosystems than NPP. While

the principle holds that catch potential is dependent on energy from

primary production, quantitative projections of catch potential are

limited by residual uncertainty on the best possible indicators of primary

production and biomass.

Assuming that the potential fish catch is proportional to NPP, the fish

catch in the North Pacific Ocean subtropical biome is projected to

increase by 26% through expansion of the biome, while catches in

the temperate and equatorial biomes may decrease by 38 and 15%,

respectively, through contraction of the biomesby 2100 relative to 2000

under the SRES A2 (RCP6.0 to 8.5) scenario (Polovina et al., 2011).

Changes in phytoplankton size structure are projected to affect fisheries

catch potential (Cheung et al., 2011), resulting in a 0 up to 75.8%

decrease in the potential catch of large fishes in the central North Pacific

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458

Chapter 6 Ocean Systems

Latitude

Depth

limate-dependent distribution

Local

xtinctions

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(

a) (b)

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Change in maximum body weight

Country A

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(e)

Invasion

hange in:

catch potential (d)

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> 50%

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no data

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eeper

00 5 10 15−10 −5

m per decade

eep

Change in maximum catch potential

Figure 6-14 | Climate change effects on the biogeography, body size, and ﬁsheries’ catch potential of marine ﬁshes and invertebrates. (a) Shifts in distribution range and

reduction in body size of exploited ﬁsh driven by projected warming, oxygen depletion, and sea ice retreat (cf. Figure 6-7). Whenever the shift in distribution does not fully

compensate for warming and hypoxia, the result will be a decrease in body size. Shifts in (b) latitudinal and (c) depth distribution of 610 exploited demersal ﬁshes are projected

to have a median (central line of the box) of 31 km per decade and 3.3 m per decade, respectively, with variation between species (box boundary: 25th and 75th percentiles)

from 1991–2010 to 2041–2060 under the SRES A2 (between RCP6.0 and 8.5) scenario (Cheung et al., 2011, 2013b). (d) Combining species’ range shifts with projected changes

in net primary production leads to a projected global redistribution of maximum catch potential. (Analysis includes approximately 1000 species of exploited ﬁshes and

invertebrates, under warming by 2°C according to SRES A1B (≈RCP6.0), comparing the 10-year averages 2001–2010 and 2051–2060; redrawn from Cheung et al., 2010.). (e)

Changes in species distribution and individual growth are projected to lead to reduced maximum body size of ﬁsh communities at a certain site. The analysis includes 610 species

of marine ﬁshes, from 1991–2010 to 2041–2060 under SRES A2 (approximately RCP6.0 to 8.5; Cheung et al., 2013b), without analysis of potential impacts of overﬁshing or

ocean acidiﬁcation. Key assumptions of the projections are that current distribution ranges reﬂect the preferences and tolerances of species for temperature and other environ-

mental conditions and that these preferences and tolerances do not change over time. Catch potential is determined by species range and net primary production. Growth and

maximum body size of ﬁshes are a function of temperature and ambient oxygen level.

459

Ocean Systems Chapter 6

nd increases of up to 43% in the California Current region over the

21st century under the SRES A2 scenario (Woodworth-Jefcoats et al.,

2013). Globally, climate change is projected to cause a large-scale

redistribution of global catch potential, with an average 30 to 70%

increase in yield at high latitudes and up to 89% in some regions, after

2°C warming from preindustrial periods following SRES A1B (≈RCP6.0)

(Cheung et al., 2010; Blanchard et al., 2012; see also Figure 6-14).

Redistribution between areas, with average catch potential remaining

unchanged, will occur at mid latitudes. A 40 to 60% drop will occur in

the tropics and in Antarctica by the 2050s relative to the 2000s (medium

confidence for direction of trends in fisheries yields, low confidence for

the magnitude of change). This highlights high vulnerabilities in the

economies of tropical coastal countries (Allison et al., 2009; see also

Section 6.4).

Fisheries targeting specific species may show more complex responses

to climate change. For example, driven by changes in temperature and

primary production, catches of skipjack and bigeye tuna in the south

Pacific are projected to increase by 2035 relative to 1980–2000 under

the SRES B1 and A2 scenario, but for 2100, skipjack tuna catch is

projected to decrease under the A2 scenario, while bigyeye tuna catch

decreases under both A2 and B1 scenarios (Lehodey et al., 2011).

Regionally, tuna catches in the Western Pacific are projected to decrease,

while those in the Eastern Pacific will increase (Lehodey et al., 2011).

Mollusk fisheries under ocean acidification is discussed under Section

6.4.1.

Identifying responses to climate change is complicated by species

interactions and multiple stressors. Major marine habitats and biodiversity

hotspots are projected to encounter cumulative impact from changes

in temperature, pH, oxygen, and primary productivity by the end of the

21st century (RCP4.5 and 8.5) (Mora et al., 2013). Acidification and

hypoxia will reduce maximum catch potential over 50 years from about

2000 onward in both the North Atlantic and Northeast Pacific

(Ainsworth et al., 2011; Cheung et al., 2011). Changes in O

content as

well as warming will drive a global decrease of community-averaged

maximum body size of 14 to 24% of exploited demersal marine fishes

by 2050 relative to the 2000s under the SRES A2 (RCP6.0 to 8.5)

scenario (Cheunget al., 2013b; see also Figure 6-14). The decrease in

maximum body size may affect natural mortality rates and trophic

interactions, and reduce yield-per-recruit and thus potential catch.

Responses of exploited marine species and their fisheries may interact

with other human stressors such as overfishing, exacerbating their

impacts (e.g., Lindegren et al., 2010; Ainsworth et al., 2011). Through

species shifts climate change may also cause overlap of habitats of

species targeted by fishing with habitat of threatened species, potentially

increasing the chances of the latter of being caught as bycatch (Jones

et al., 2013). Moreover, differences in vulnerability and adaptive capacity

of species to changing environmental and ecosystem conditions will

affect the responses of fisheries to climate change (e.g., Le Borgne et

al., 2011; Griffith et al., 2011).

The complex and nonlinear interactions and responses of both biophysical

and socioeconomic systems to climate change may lead to changes that

have a low probability of occurrence based on empirical data (Doak et

al., 2008). The risk of such low-probability but potentially high-impact

events may be underestimated in existing model projections (Williams

nd Jackson, 2007; Lindenmayer et al., 2010). Projected changes in the

distribution and production potential of fisheries resources are expected

to affect economics, human livelihood, and food security (Allison et al.,

2009; Sumaila and Cheung, 2010; low confidence in the magnitude and

direction of the projected socioeconomic impacts).

.5.4. Conclusions

Modeling projects that the distribution of invertebrates, fishes, and

some marine mammals, birds, and reptiles will shift further under most

emission scenarios, with rates and directions of shifts consistent with

those observed in the last century (high confidence; Sections 6.3.1-7).

These projections are valid for those species that adapt not at all or

incompletely to warmer temperatures and the associated ecosystem

changes, as indicated by present trends (Section 6.3.1; Box CC-MB). For

non-adapting species rates of shift will thus increase with increasing

rates of warming and higher emission scenarios (high confidence),

unless the shift is blocked by geographic or other barriers (e.g., light

regime; Figure 6-7). The average shift in distribution will continue to be

poleward at large spatial scales (high confidence; Section 6.5.2; Box

CC-MB). Species richness and the abundance of warm-water species

will increase at high latitudes (high confidence) and decrease in the

tropics (medium confidence; Section 6.5.2). Projections for individual

species and populations are more variable and sensitive to model

parameters.

Maximum fisheries catch potential is projected to increase at high

and decrease at low latitudes by 2050 under SRES B1 (≈RCP4.5) and

A1B (≈RCP6.0) climate scenarios (medium confidence; Section 6.5.3).

Quantifying such projections is constrained by uncertainties in projected

primary production rates (Sections 6.3.4, 6.5.1), biological responses

such as species interactions (Section 6.3.6), and in projected effects of

multiple climate drivers and human activities (low confidence; Section

6.3.5).

Models that integrate climate and ocean changes with biological

responses and interactions, and with current human activities, have led

to agreement on species and food web responses to climate change

(Section 6.5.3). However, most of these models do not include trophic

interactions. They insufficiently consider physiological principles and

none include evolutionary adaptations that affect responses of biota to

physical and chemical changes.

Projections of ocean biogeochemistry represent the open oceans rather

well, but coastal and shelf regions only poorly. From a global perspective,

open ocean NPP will decrease moderately by 2100 under both medium

(SRES B1 or RCP4.5) and high emission scenarios (medium confidence;

A2 or RCP6.0 to 8.5; Sections 6.3.4, 6.5.1), paralleled by an increase in

NPP at high latitudes and a decrease in the tropics (medium confidence;

Sections 6.3.4, 6.5.1; Box CC-PP).

Overall, the projected responses of marine organisms and ecosystems

to climate change include changes in primary productivity (medium

confidence), species’ life history (medium confidence), distribution,

abundance, and diversity across marine food webs (high confidence) in

a time frame of 20 to 80 years from 2010, with substantially larger

460

Chapter 6 Ocean Systems

ong-term (end of 21st century) responses under high emission scenarios

(high confidence). These changes will be largest under business-as-usual

scenarios (RCP8.5) and increase the vulnerability of human societies,

y affecting income, employment, and food security through their

effects on fisheries, tourism, and regulatory services such as coastal

protection (medium confidence; Section 6.4.1.3; Box CC-CR).

tlantic Cod (AC)

Sardines and Anchovies, Japan Sea (SAJ)

Paciﬁc Salmon (PS)

elpout, Wadden Sea (EWS)

anded Morwong (BM)

arine Air Breathers (MAB)

acroorganism Effects (MAE, animals and plants)

lankton Phenology (PP)

olar Organisms (PO)

eef-building Warm Water Corals (RWC)

id-water Fishes (MWF)

yster Effects (EO)h

alcifying Organisms (CAL)

10 11

75b

1110

Low Very low Low Medium

Conﬁdence in attribution

High

Very high

Very low Low Medium

Conﬁdence in detection and/or projection

High Very high

Very low

Medium

Conﬁdence in attribution

High Very high

Very low Low Medium

Conﬁdence in detection and/or projection

High Very high

Detection

Projection

Detection and projection have the same levels of conﬁdence

Oxygen and Capacity Limited Thermal Tolerance (OCLTT)

Geological Record (GR, observations)

Global Net Primary Production (gNPP)

Synergistic Effects (SE)

Fishery Catch Potential (FCP, species shifts)

Community Composition (CC, under TE, HE, OAE)

Biogeochemical Processes (BG)

Abundance (AB)

Ocean Acidiﬁcation Effects (OAE)

High-latitude Net Primary Production (hNPP)

Hypoxia Effects (HE)

Temperature Effects (TE)

Species Richness (SR, Fish)

Speciﬁc examples

Broad categories

4 Ecosystem Services (ES)

5b Fishery Catch Potential (FCP, changing NPP)

Harmful Algae Blooms (HAB)

Detection

Projection

Detection and projection have the same levels of conﬁdence

Figure 6-15 | Overview of the levels of conﬁdence in detection, as well as in projection, of climate change effects on ocean systems, in relation to the levels of conﬁdence in

attributing these effects to the respective climate forcings. Case studies, processes, and concepts relevant in assessing the effects of climate change are represented by their

acronyms in both text and ﬁgure. While conﬁdence in the presence of effects is often high, the direct attribution to one driver in ﬁeld experiments is difﬁcult, as drivers are often

highly correlated with each other (e.g., warming with changes in stratiﬁcation, hence reduced nutrient supply). Some climate change impacts have been condensed into broad

categories to avoid overpopulating the ﬁgures (e.g., Bio-Geochemical processes, BG). Note that the term “attribution” is used for both present-day detections in the ﬁeld and

future projections, the latter including qualitative and quantitative extrapolations and simulations of future conditions from fundamental principles, experiments, and models. Firm

knowledge from experiments (ﬁeld, laboratory, and modeling) simulating future conditions enhances the respective conﬁdence levels to those for detection or projection. The

empirical observations resulting from those experiments are directly attributable to the respective drivers. Conﬁdence in attribution is enhanced if these experiments identify the

underlying mechanisms and their responses. See text for the discussion of depicted examples and categories. Conﬁdence assignments focus on the nature and size of effects, not

on model capacity to quantify their magnitude reliably.

461

Ocean Systems Chapter 6

6.6. Chapter Conclusions and Key Uncertainties

This section provides an overview of confidence levels in the detection

and projection of climate change effects on ocean systems, and of

onfidence levels in their attribution to different forcings. It distinguishes

between effects previously observed and those projected, and considers

confidence in the knowledge of underlying principles as discussed in

this chapter. While the anthropogenic signal is conspicuous in the oceans

(Section 6.1.1), clear attribution to anthropogenic influences on climate

is not always possible in individual case studies, owing to the inherent

variability of the system (Figure 6-15; acronyms of relevant processes,

capitalized, link between text and figure).

Present-day observations and those from the Geological Record (GR;

Figure 6-15) show similar signs of response to environmental changes, for

example, warming at high CO

levels, and similar ecological consequences

in the ocean (robust evidence, medium agreement; medium confidence).

However, the ongoing rate of anthropogenic CO

release and hence

ocean acidification is unprecedented in the last 65 Ma (high confidence)

and probably the last 300 Ma (Section 6.1.2).

6.6.1. Key Risks Related to Climate Change:

Constraints on Ecosystem Services

Empirical studies provide evidence that climate change has impacted

marine ecosystems (high confidence; FAQ 6.4; Table 6-6) and has caused

changes in provisioning, regulating, and supportive Ecosystem Services

(ES; medium confidence). Climate change may also have affected cultural

services (limited evidence, medium agreement) but attribution of

impacts to these services remains a challenge (low confidence), owing

to the intrinsic difficulties of assessing these services, the lack of long

time-series data, and confounding human impacts. In light of available

understanding of cause and effect of climate change impacts on marine

ecosystems (high confidence), future climate change will affect some

ecosystem services (high confidence in projection, medium confidence in

attribution). Projected changes in the availability of marine resources and

ecosystem services are expected to affect economics, human livelihood,

and food security. Vulnerability is highest for the national economies of

tropical coastal countries (high confidence).

6.6.1.1. Redistribution and Constraints on

Microbial Functions and Primary Productivity

Laboratory and mesocosm studies have identified various microbially

mediated processes responding to climate-induced changes in light,

nutrient supply, temperature, CO

, and hypoxia (high confidence). Such

processes include nitrogen fixation and the nitrogen cycle, carbon

sequestration and export production, calcification, respiration, O

production, climate-feedback by dimethylsulfide (DMS) production, and

nutrient recycling. However, changes in these Bio-Geochemical processes

(BG) in the field are difficult to detect, project, and attribute to climate

change (low confidence; Sections 6.3.1-5).

The trends in net primary production recently reported for much of the

low-latitude ocean using satellite observations differ considerably from

hose few long-term direct estimates of NPP at oceanic time series sites

(Sections 6.1.2, 6.3.4). Increased NPP at high latitudes (hNPP, detected

and attributable to climate change with medium confidence; Section

6.3.4; Box CC-PP) are indicated by satellite images (medium confidence)

and due to reduction and thinning of sea ice. Trends in NPP will be

strengthened with further warming (medium confidence). Modeling

projects that global NPP (gNPP) will decrease by 2100 under RCP

scenarios (medium confidence; Section 6.5.1; Box CC-PP).

6.6.1.2. Warming-Induced Species Redistribution,

Loss of Biodiversity, and Fisheries Catch Potential

Long-term observations show variability in oceanographic conditions

with a key role of temperature and changing oceanographic regimes

causing observed changes in ecosystem structure and fish stocks (very

high confidence; cf. Section 30.7.1.1). Temperature Effects (TE) reflect the

differential specialization of all life forms in limited ambient temperature

ranges (very high confidence). Temperature exerts strong MAcroorganism

Effects (MAE), that is, on animals and plants. Warming is presently

causing and will cause species displacements and largely poleward

shifts in biogeographic distribution of zooplankton and fishes, paralleled

by altered seasonal activity, species abundance, migration, and body

size (high to very high confidence; Section 6.3.1), and leading to shifts

in Community Composition (CC; high confidence; Box 6-1). Causes and

effects are understood for fishes and most invertebrates via their

Oxygen and Capacity Limited Thermal Tolerance (OCLTT; robust

evidence, medium agreement; high confidence; Section 6.3.1). Such

knowledge supports projections into the future (high confidence;

Section 6.5), which are influenced by the limited potential of organisms

to adapt. Alterations in species ABundance (AB) result when organisms

encounter shifting mean and extreme temperatures (high confidence

in detection and attribution). Such trends will be exacerbated during

future warming (high confidence; Section 6.5.1).

Among prominent examples, warming has caused and will cause

northward shift and expansion of the geographic distribution of North

Atlantic Cod (AC; high confidence in detection or projection, medium

confidence in detection or projection and attribution; Section 6.3.1) and

shifting growth patterns in relation to the distribution of Banded

Morwong around New Zealand (BM; high confidence in detection or

projection, medium confidence in detection or projection and attribution).

Warming has shifted dominant species from Sardines to Anchovies in the

Sea of Japan (SAJ; medium confidence in detection, medium confidence

in detection and attribution; Sections 6.3.1, 6.3.6). Warming extremes

have reduced and will further reduce the abundance of Eelpout in the

Wadden Sea (EWS; high confidence in detection or projection, high

confidence in detection or projection and attribution; Section 6.3.1).

Extreme warming events increase mortalities of Pacific Salmon during

spawning migrations (PS; high confidence in detection, high confidence

in detection and attribution; Section 6.3.1) in Fraser River, Canada. At

temperate and high latitudes, communities display increasing fish Species

Richness (SR) resulting from latitudinal shifts of species and attributed

to warming and loss of sea ice, although the relative contributions of

regional climate variation and long-term global trends have not been

quantified (high confidence in detection, medium confidence in detection

and attribution; Sections 6.3.1, 6.5.2). Latitudinal species shifts are

462

Chapter 6 Ocean Systems

Key risk Adaptation issues & prospects

Climatic

drivers

Risk & potential for

adaptation

Timeframe

Risks to ecosystems and adaptation options

Present

°C

4°C

ery

low

Very

high

edium

Present

2°C

4°C

Very

ery

high

Medium

Present

2°C

4°C

Very

ery

high

Medium

Present

2°C

4°C

Very

low

Very

high

Medium

Present

2°C

4°C

Very

low

Very

high

Medium

Present

2°C

4°C

Very

low

Very

high

Medium

Table 6-6 | Coastal and oceanic key risks from climate change and the potential for risk reduction through mitigation and adaptation. Key risks are identiﬁed based on

ssessment of the literature and expert judgments made by authors of the various WGII AR5 chapters, with supporting evaluation of evidence and agreement in the referenced

chapter sections. Each key risk is characterized as very low, low, medium, high, or very high. Risk levels are presented for the near-term era of committed climate change (here,

or 2030–2040), in which projected levels of global mean temperature increase do not diverge substantially across emissions scenarios. Risk levels are also presented for the

longer-term era of climate options (here, for 2080–2100), for global mean temperature increase of 2°C and 4°C above pre-industrial levels. For each time frame, risk levels are

estimated for the current state of adaptation and for a hypothetical highly adapted state. As the assessment considers potential impacts on different physical, biological, and

human systems, risk levels should not necessarily be used to evaluate relative risk across key risks. Relevant climate variables are indicated by symbols. Acronyms for oceans

sub-regions are as follows: HLSBS = High-Latitude Spring Bloom Systems; EUS = Equatorial Upwelling Systems; SES = Semi-Enclosed Seas; CBS = Coastal Boundary Systems;

EBUE = Eastern Boundary Upwelling Ecosystems; STG = Sub-Tropical Gyres, DS = Deep Sea (>1000 m).

Near term

(2030 – 2040)

Long term

(

2080 – 2100)

ear term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

hanges in ecosystem productivity

ssociated with the redistribution and

loss of net primary productivity in open

oceans. (medium conﬁdence)

[

6.5.1, 6.3.4, 30.5.1-2, Box CC-PP]

daptation options are limited to the translocation of industrial ﬁshing activities

ue to regional decreases (low latitude) versus increases (high latitude) in

productivity, or to the expansion of aquaculture.

Distributional shift in ﬁsh and invertebrate

species, fall in ﬁsheries catch potential at

ow latitudes, e.g., in EUS, CBS, and STG

regions. (high conﬁdence)

[6.3.1, 6.5.2-3, 30.5.1-4, 30.6.2,

Box CC-MB]

Evolutionary adaptation potential of ﬁsh and invertebrate species to warming is limited

as indicated by their changes in distribution to maintain temperatures. Human adaptation

options involve the large-scale translocation of industrial ﬁshing activities following the

regional decreases (low latitude) versus (possibly transient) increases (high latitude) in

catch potential as well as deploying ﬂexible management that can react to variability and

change. Further options include improving ﬁsh resilience to thermal stress by reducing

other stressors such as pollution and eutrophication, the expansion of sustainable

aquaculture and development of alternative livelihoods in some regions.

High mortalities and loss of habitat to

larger fauna including commercial species

due to hypoxia expansion and effects.

(high conﬁdence)

[6.3.3, 30.5.3-5]

Human adaptation options involve the large-scale translocation of industrial

ﬁshing activities as a consequence of the hypoxia-induced decreases in

biodiversity and ﬁsheries catch of pelagic ﬁsh and squid. Special ﬁsheries may

beneﬁt (Humboldt squid). Reducing the amount of organic carbon running off

coastlines by controlling nutrients and pollution running off agricultural areas can

reduce microbial activity and consequently limit the extent of the oxygen

drawdown and the formation of coastal dead zones.

Ocean acidiﬁcation: Reduced growth and

survival of commercially valuable shellﬁsh

and other calciﬁers, e.g., reef building

corals, calcareous red algae.

(high conﬁdence)

[5.3.3.5, 6.1.1, 6.3.2, 6.4.1.1, 30.3.2.2,

Box CC-OA]

Evidence for differential resistance and evolutionary adaptation of some species

exists but is likely limited by the CO

concentrations and high temperatures

reached; adaptation options include the shift to exploiting more resilient species

or the protection of habitats with low natural CO

levels, as well as the reduction

of other stresses, mainly pollution and limiting pressures from tourism and ﬁshing.

Reduced biodiversity, ﬁsheries abundance

and coastal protection by coral reefs due

to heat-induced mass coral bleaching and

mortality increases, exacerbated by ocean

acidiﬁcation, e.g., in CBS, SES, and STG

regions. (high conﬁdence)

[5.4.2.4, 6.3.1, 6.4.2, 30.3.1.1, 30.3.2.2,

30.5.3-6, Box CC-CR]

Evidence of rapid evolution by corals is very limited or nonexistent. Some corals

may migrate to higher latitudes. However, the movement of entire reef systems is

unlikely given estimates that they need to move at the speed of 10 – 20 km yr

–1

Human adaptation options are limited to reducing other stresses, mainly

enhancing water quality and limiting pressures from tourism and ﬁshing. This

option will delay the impacts of climate change by a few decades but is likely to

disappear as thermal stress increases.

Coastal inundation and habitat loss due

to sea level rise, extreme events, changes

in precipitation, and reduced ecological

resilience, e.g., in CBS and STG

subregions. (medium to high conﬁdence)

[5.4.2.3-7, 5.5.2, 5.5.4, 30.5.6, Box

CC-CR]

Options to maintain ecosystem integrity are limited to the reduction of other

stresses, mainly pollution and limiting pressures from tourism, ﬁshing, physical

destruction, and unsustainable aquaculture; reducing deforestation and increasing

reforestation of river catchments and coastal areas to retain sediments and

nutrients; increased mangrove, coral reef, and seagrass protection and restoration

to protect numerous ecosystem goods and services such as coastal protection,

tourist value, and ﬁsh habitat.

Damaging

yclone

Ocean

cidiﬁcation

Precipitation

Climate-related drivers of impacts

Warming

rend

Extreme

recipitation

Extreme

emperature

Sea

evel

Level of risk & potential for adaptation

otential for additional adaptation

o reduce risk

isk level with

urrent adaptation

isk level with

igh adaptation

Hypoxia

Continued next page

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Present

2°C

4°C

Very

low

Very

high

Medium

Marine biodiversity loss with high rate of

climate change. (medium conﬁdence)

[6.3.1-3, 6.4.1.2-3, Table 30.4, Box

CC-MB]

Adaptation options are limited to the reduction of other stresses, mainly to

reducing pollution and to limiting pressures from tourism and ﬁshing.

463

Ocean Systems Chapter 6

projected to continue in the 21st century under all IPCC emission

scenarios (high confidence; Sections 6.3.1, 6.3.5, 6.3.7, 6.4.1, 6.5.2).

Climate-induced regime shifts and regional changes in Plankton

Phenology (PP; medium confidence) have caused and will cause changes

in food composition and availability to animals. Species shifts and

changing species composition lead to changes in Fishery Catch Potential

(FCP; high confidence; 5a in Figure 6-15), partly attributable to climate

change (high confidence) and to sustained fishing pressure (Section

6.5.3). Fisheries Catch Potentials (FCP) will be redistributed, decrease

at low latitudes, and increase at high latitudes (high confidence; 5a in

Figure 6-15). These trends will possibly be strengthened by the projected

decrease in NPP at low latitudes and increase in NPP at high latitudes

(medium confidence; Sections 6.5.2-3; 5b in Figure 6-15). Polar Organisms

(PO) that are unable to migrate to cooler waters, and to acclimatize or

to adapt to warming, will become marginalized, contributing to the

projected high species turnover in polar areas (high confidence; Sections

6.3.1, 6.5.2).

Detected effects on Marine Air Breathers (MAB: mammals, seabirds, and

reptiles) include changing abundances and phenology, shifts in species

distribution, and in sea turtle sex ratios (high confidence), all of which

are partly attributed to climate change (high confidence). However, few

effects are directly linked to climate drivers (e.g., temperature-driven

turtle sex ratio); most effects are due to shifts in habitat structure (e.g.,

loss of sea ice), changing availability of prey organisms, or changes in

Present

°C

4°C

Very

ery

igh

Medium

Present

2°C

4°C

ery

low

Very

high

edium

Redistribution of catch potential of large

elagic-highly migratory ﬁsh resources, such as

tropical Paciﬁc tuna ﬁsheries. (high conﬁdence)

[

6.3.1, 6.4.3, Table 30.4]

International ﬁsheries agreements and instruments, such as the tuna

ommissions, may have limited success in establishing sustainable

ﬁsheries yields.

Variability of small pelagic ﬁshes in Eastern

Boundary Upwelling systems is becoming more

extreme at interannual to multi-decadal scales,

making industry and management decisions more

uncertain. (medium conﬁdence)

[6.3.2, 6.3.3, 30.5.5, Box CC-UP]

Development of new and speciﬁc management tools and models

may have limited success to sustain yields. Reduction in ﬁshing

intensity increases resilience of the ﬁsheries.

Present

2°C

4°C

Very

low

Very

high

Medium

Present

2°C

4°C

Very

low

Very

high

Medium

Decrease in catch and species diversity of ﬁsheries in

tropical coral reefs, exacerbated by interactions with

other human drivers such as eutrophication and

habitat destruction. (high conﬁdence)

[6.4.1, 30.5.3-4, 30.5.6, Table 30-4, Box CC-CR]

Restoration of overexploited ﬁsheries and reduction of other

stressors on coral reefs delay ecosystem changes. Human adaptation

includes the usage of alternative livelihoods and food sources (e.g.,

coastal aquaculture).

Current spatial management units, especially the

MPAs, may fail in the future due to shifts in species

distribution and community structure.

(high conﬁdence)

[6.3.1, 6.4.2.1, 30.5.1, Box CC-MB]

Continuous revision and shifts of MPA borders, and of MPA goals

and performance.

Present

2°C

4°C

Very

low

ery

high

edium

Present

2°C

4°C

Very

low

Very

high

Medium

Decreased production of global shellﬁsh ﬁsheries.

(

high conﬁdence)

[6.3.2, 6.3.5, 6.4.1.1, 30.5.5, 30.6.2.1, Box CC-OA]

Effective shift to alternative livelihoods, changes in food

onsumption patterns, and adjustment of (global) markets.

lobal redistribution and decrease of low-latitude

ﬁsheries yields are paralleled by a global trend to

atches having smaller ﬁshes. (medium conﬁdence)

[6.3.1, 6.4.1, 6.5.3, 30.5.4, 30.5.6, 30.6.2]

ncreasing coastal poverty at low latitudes as ﬁsheries becomes

smaller – partially compensated by the growth of aquaculture and

arine spatial planning, as well as enhanced industrialized ﬁshing

efforts.

Near term

(2030 – 2040)

ong term

(

2080 – 2100)

ear term

(

2030 – 2040)

ong term

(

2080 – 2100)

ear term

(

2030 – 2040)

ong term

(

2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Key risk Adaptation issues & prospects

Climatic

drivers

isk & potential for

adaptation

Timeframe

Risks to ﬁsheries

Continued next page

Table 6-6 (continued)

Continued next page

464

Chapter 6 Ocean Systems

foraging efficiency, in both mammals (polar bears, walruses) and birds

(penguins, albatrosses). Such trends will be exacerbated by future

warming (high confidence; Sections 6.3.7, 6.5.2).

6.6.1.3. Expanding Hypoxia Affecting Marine Resources

Hypoxic zones in marine sediments and pelagic OMZs will continue to

expand in the future, owing to climate-induced warming trends (Section

6.1.1). Local and regional Hypoxia Effects (HE) have been observed

(medium confidence) and will be exacerbated in the future (high

confidence; Section 6.3.3) causing habitat loss for groundfishes and

pelagic predators and affecting the distribution of key zooplankton and

nekton species (medium confidence). Progressive hypoxia is causing

shifts in community composition toward hypoxia-tolerant species,

excluding calcifiers due to elevated pCO

(high confidence), benefiting

specialized microbes, and leading to reduced biodiversity and the loss

of higher life forms (high confidence; Section 6.3.3). Loss of deep habitat

and biomass of Mid-Water Fishes (MWF; Section 6.3.3; medium

confidence in detection) off California is also attributed to hypoxia

(medium confidence). These trends will continue into the future (medium

confidence).

6.6.1.4. Constraints on Marine Calcifiers and Associated Fisheries

and Aquaculture due to Ocean Acidification

Ocean acidification will exert negative effects on species and whole

ecosystems and their services, especially those relying on carbonate

structures such as warm-water coral reefs (high confidence; cf. Section

30.7.1.2). Presently, only a small number of field observations have

detected Ocean Acidification Effects (OAE; medium confidence), but

experiments and natural analogs support reliable but qualitative

projections and attribution (high confidence). A specific glimpse into

the future of anthropogenic OA is provided by negative Effects of

upwelled CO

-rich waters on Pacific Oysters (EO) introduced to

aquaculture along the North American west coast (high confidence in

detection, low confidence in attribution to anthropogenic causes).

Findings in experimental laboratory and field studies as well as at

natural analogs support attribution of projected effects to future CO

Key risk Adaptation issues & prospects

Climatic

drivers

isk & potential for

adaptation

Timeframe

Risks to humans and infrastructure

resent

2°C

4°C

Very

low

ery

igh

edium

Present

°C

4°C

Very

low

Very

igh

Medium

Impacts due to increased frequency of

armful algal blooms (medium conﬁdence)

[6.4.2.3, 30.6.3]

Adaptation options include improved monitoring and early warning system,

eduction of stresses favoring harmful algal blooms, mainly pollution and

utrophication, as well as the avoidance of contaminated areas and ﬁsheries

products.

Impacts on marine resources threatening

regional security as territorial disputes and

food security challenges increase

(limited evidence, medium agreement)

[AR5 SREX, 30.6.5, 30.7.2, 12.4-6, 29.3]

Decrease in marine resources, movements of ﬁsh stocks and opening of new

seaways , and impacts of extreme events coupled with increasing populations

will increase the potential for conﬂict in some regions, drive potential

migration of people, and increase humanitarian crises.

Present

2°C

4°C

Very

low

Very

igh

Medium

Impacts on shipping and infrastructure for

energy and mineral extraction increases as

storm intensity and wave height increase

in some regions (e.g., high latitudes)

(high conﬁdence)

[AR5 SREX, 30.6.2.3-4, 30.6.5, 29.3]

Adaptation options are to limit activities to particular times of the year and/or

develop strategies to decrease the vulnerability of structures and operations.

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Near term

(2030 – 2040)

Long term

(2080 – 2100)

Table 6-6 (continued)

Present

2°C

4°C

Very

low

Very

igh

edium

Reduced livelihoods and increased poverty.

(medium conﬁdence)

[6.4.1-2, 30.6.2, 30.6.5]

Human adaptation options involve the large-scale translocation of industrial

ﬁshing activities following the regional decreases (low latitude) versus increases

(high latitude) in catch potential and shifts in biodiversity. Artisanal local

ﬁsheries are extremely limited in their adaptation options by available ﬁnancial

resources and technical capacities, except for their potential shift to other

species of interest.

Near term

(2030 – 2040)

ong term

(2080 – 2100)

Present

2°C

4°C

Very

high

Medium

Coastal socioeconomic security.

(high conﬁdence)

[

5.5.2, 5.5.4, 30.6.5, 30.7.1, Table 30-4]

Human adaptation options involve (1) protection using coastal defences (e.g.

eawalls) and soft measures (e.g., mangrove replanting and enhancing coral

rowth); (2) accommodation to allow continued occupation of coastal areas by

making changes to human activities and infrastructure; and (3) managed retreat

as a last viable option. Options vary from large-scale engineering works to

maller scale community projects. Options are available under the more

traditional CZM (coastal zone management) framework but increasingly under

DRR (disaster risk reduction) and CCA (climate change adaptation) frameworks.

*High conﬁdence in existence of adaptation measures, Low conﬁdence in magnitude of risk reduction

ear term

(

2030 – 2040)

ong term

(

2080 – 2100)

465

Ocean Systems Chapter 6

oncentrations (medium confidence), with species-specific sensitivities

across phyla (high confidence). Projected effects are most harmful to

strong CALcifiers (CAL; high confidence), for example, some echinoderms,

bivalves, gastropods, warm-water corals, and crustose algae, and less

harmful to some crustaceans and, possibly, fishes. Projections from

experimental studies and observations at natural analogs indicate shifts

in Community Composition (CC) to more active animals and from

calcifiers (CAL) to non-calcifiers in all organism groups (high confidence

in both projection and attribution to increased CO

; Section 6.3.2;

Table 6-3).

6.6.1.5. Interactions of Climate-Related Drivers Exacerbating

Impacts on Organisms, Ecosystems, and Their Services

Climate change involves interactions of temperature with other climate-

related drivers and their effects (ocean acidification, hypoxia, freshening,

nutrient supply, organism shifts resulting in changing interactions

between species, changes in habitat structure, e.g., loss of sea ice).

Strong interactions with other human impacts like eutrophication, fishing,

and other forms of harvesting accelerate and amplify climate-induced

changes (high confidence; Section 6.3.5, 30.7.1.1). Harmful algal blooms

(HAB) will be stimulated by warming, nutrient fluctuations in upwelling

areas, eutrophication in coastal areas (Table 6-6), ocean acidification,

and enhanced surface stratification (medium confidence). Synergistic

Effects (SE) will be exacerbated in the future (medium confidence), but

have not yet been clearly detected and attributed in the field (low

confidence). For projected future effects, attribution of observed impacts

to such synergisms is supported by experimental evidence, especially

in animals and plants (medium confidence).

Increased bleaching and decreased calcification displayed by several

Reef-building Warm-water Corals (RWC; very high confidence) over the

last 3 decades are attributed to the ongoing warming trend, and the

associated rise in extreme temperature events and amplitudes (high

confidence; Sections 6.3.1, 30.5.6; Box CC-CR). Such trends will be

exacerbated by future warming and synergistic effects (high confidence;

cf. Section 30.5.4.2), with some amelioration by latitudinal shifts and

evolutionary adaptation (Section 6.3.1; low confidence). Ocean

acidification will have an increasing influence on reefs (high confidence),

as indicated by similar phenomena during mass extinctions in Earth history.

6.6.2. Key Uncertainties

Key uncertainties result from insufficient knowledge of ocean systems.

International organizations (both inter- and non-governmental) have

the opportunity to play a key role in coordinating research concepts and

approaches, working toward a coherent picture of climate change

effects on the global ocean. Countries around the world have limited

capacity and infrastructure to study the ocean’s response to climate

change. Long-term observational time series are especially lacking, in

both quantity and quality. Research has provided valuable insights, but

a unifying approach addressing principles across organism domains and

ecosystems is still missing. Processes investigated so far differ largely

by study organisms (plants, animals, phytoplankton, and bacteria)

and by level of organization (ecosystem, whole organism, tissue, cell,

olecular). Especially for microbes, available data are patchy and

reported trends are often in different directions, partly due to different

experimental protocols and/or over-reliance on species or strains of

microbes that are readily culturable, and hence have beenused for

decades in laboratory research. The knowledge base of climate impacts

on species, strains, or communities in the field is insufficient. Scaling from

physiological studies on individual species to ecosystem changes has

been successful in individual cases but has not been widely implemented,

for example, to shifts in species interactions or food webs. An integrated

framework of climate sensitivity at the ecosystem level that considers

multiple drivers and their interactive effects needs to be developed

further. This includes an in depth understanding of ecosystem structure

(physical and biological) and functioning, of ecosystem complexity and

species interactions, and of the resulting implications for biogeochemical

processes. For all climate drivers, especially ocean warming, acidification,

and hypoxia, studies integrating mechanistic knowledge and evolutionary

adaptation over generations are needed. Research should also cover

various climate zones and biomes. Laboratory and modeling experiments

are needed to test hypotheses building on long-term field observations

and observations at natural or paleo-analogs. Models should better

integrate observations and mechanism-based understanding, and better

project future interactions between human and natural systems in a

changing climate.

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