Life in most of the global ocean, from pole to pole and from sea surface to the abyssal depths, is already experiencing higher temperatures due to human-driven climate change. In many places, that increase may be barely measurable. In others, particularly in near-surface waters, warming has already had dramatic impacts on marine animals, plants and microbes. Due to closely linked changes in seawater chemistry, less oxygen remains available (in a process called ocean deoxygenation). Seawater contains more dissolved carbon dioxide, causing ocean acidification. Non-climatic effects of human activities are also ubiquitous, including over-fishing and pollution. Whilst these stressors and their combined effects are likely to be harmful to almost all marine organisms, food-webs and ecosystems, some are at greater risk (FAQ5.1, Figure 1). The consequences for human society can be serious unless sufficient action is taken to constrain future climate change.
Ocean oxygen changes are also affected by climate variability on interannual and decadal timescales, especially for the tropical ocean OMZs (Deutsch et al., 2011). ENSO variability in particular affects the thermocline structure, which then alongside changes in circulation modulates oxygen solubility and respiratory demand in this region (Ito and Deutsch, 2013; Eddebbar et al., 2017). These drivers may then be combined with modifications to overturning and ventilation of OMZs by lateral jets and equatorial current intensity (Duteil et al., 2014). Centennial scale studies based on isotope proxies for low oxygen regions have demonstrated fluctuations in OMZ extent linked to decadal changes in tropical trade winds that affects interior ocean respiratory oxygen demand, which implies that it will be difficult to attribute recent changes in the Pacific OMZ to anthropogenic forcing alone (Deutsch et al., 2015). Parallel work based on oxygen observations (Llanillo et al., 2013), as well as modelling (Duteil et al., 2018) supports the importance of decadal scale variability in the eastern tropical Pacific OMZ. There is some evidence for the potential of a modulating impact on tropical Pacific oxygen at interannual timescales from atmospheric deposition of nitrogen and iron (Ito et al., 2016; Yang and Gruber, 2016).
Detailed analyses of the interplay between different drivers of NPP, including temperature, light, nutrient levels and grazing from a subset of CMIP5 models, reveals a complex interplay with a strong latitudinal dependence (Laufkötter et al., 2015330) summarised in Figure 5.12. Warming acts to enhance growth, most notably at lower latitudes, while light conditions are also predicted to improve, mostly at the poles. Nutrient limitation shows a much more complex response across models, but tends to increase in the tropics and northern high latitudes, with little change in the Southern Ocean. Taken together there is a tendency for reduced growth rates across the entire ocean, but there is a large amount of inter-model variability. The changes in growth are allied to a consistent increase in the grazing loss of biomass to upper trophic levels. Since AR5, we have an increasing body of literature concerning role of biological feedbacks, especially due to interactions between organisms, specific physiological responses and from upper trophic levels on nutrient concentrations, linked to variable food quality (Kwiatkowski et al., 2018331), resource recycling (Boyd et al., 2015a332; Tagliabue et al., 2017333) and interactions between organisms (Lima-Mendez et al., 2015334), but their role in shaping the response of NPP to climate change remains a major unknown. Lastly, modelling work suggests that the increasing deposition of anthropogenic aerosols (supplying N and Fe) stimulates biological activity (Wang et al., 2015b335) and may compensate for warming driven reductions in primary productivity (Wang et al., 2015b336), but these effects do not form part of the CMIP5 projections assessed here.
WGII AR5 concluded that the vulnerability of most organisms to warming is set by their physiology, which defines their limited temperature ranges and thermal sensitivity (Pörtner et al., 2014385). Although different hypotheses have been proposed since AR5 to explain the mechanism linking temperature sensitivity of marine organisms and their physiological tolerances (Schulte, 2015386; Pörtner et al., 2017387; Somero et al., 2017388), evidence from physiological experiments and observations from paleo- and contemporary periods continue to support the conclusion from AR5 on the impacts of temperature change beyond thermal tolerance ranges on biological functions such as metabolism, growth and reproduction (Payne et al., 2016389; Pörtner and Gutt, 2016390; Gunderson et al., 2017391), contributing to changes in biogeography and community structure (Beaugrand et al., 2015392; Stuart-Smith et al., 2015393) (high agreement, high confidence). Comparison of biota across land and ocean suggests that marine species are generally inhabiting environment that is closer to their upper temperature limits, explaining the substantially higher rate of local extirpation related to warming relative to those on land (Pinsky et al., 2019394). Hypoxia and acidification can also limit the temperature ranges of organisms and exacerbate their sensitivity to warming (Mackenzie et al., 2014395; Rosas-Navarro et al., 2016396; Pörtner et al., 2017397), although interactions vary strongly between species and biological processes (Gobler and Baumann, 2016398; Lefevre, 2016399).
WGII AR5 concludes that multiple climatic hazards from ocean acidification, hypoxia and decrease in nutrient and food supplies pose risks to marine ecosystems, and the risk can be elevated when combined with warming (Riebesell and Gattuso, 2014512; Gattuso et al., 2015513). In a recent meta-analysis of 632 published experiments, primary production by temperate non-calcifying plankton increases with elevated temperature and CO2, whereas tropical plankton decreases productivity because of acidification (Nagelkerken and Connell, 2015514). Also, temperature increases consumption and metabolic rates of herbivores but not secondary production; the latter decreases with acidification in calcifying and non-calcifying species. These effects together create a mismatch with carnivores whose metabolic and foraging costs increase with temperature (Nagelkerken and Connell, 2015515). Warming may also exacerbate the effects of ocean acidification on the rate of photosynthesis in phytoplankton (Lefevre, 2016516). There is some, but limited, reports of observed impacts on calcified pelagic organisms that are attributed to secular trend in ocean acidification and warming (Harvey et al., 2013517; Kroeker et al., 2013518; Nagelkerken et al., 2015519; Boyd et al., 2016520). For example, Rivero-Calle et al. (2015) reported, using CPR archives, that stocks of coccolithophores (a group of phytoplankton that forms calcium carbonate plateles) have increased by 2% to over 20% in the north Atlantic over the last five decades, and that this increase is linked to synergistic effects of increasing anthropogenic CO2 and rising temperatures, as supported by their statistical analysis and a number of experimental studies. Most of the available evidence supports that ocean acidification and hypoxia can act additively or synergistically between each other and with temperature across different groups of biota (Figure 5.13). Limitation of nutrient and food availability and predation pressures can further increase the sensitivity of organismal groups to climate change in specific ecosystems (Riebesell et al., 2017494). Climate change also affects organisms indirectly through the impacts on competitiveness between organisms that favour those that are more adaptive to the changing environmental conditions (Alguero-Muniz et al., 2017495) and changes in trophic interactions (Seebacher et al., 2014496). Overall, direct in situ observations and laboratory experiments show that there are significant responses to the multiple stressors of warming, ocean acidification and low oxygen on phytoplankton, zooplankton and fishes and that these responses can be additive or synergistic (high confidence, Figure 5.13).
Changes in fish catches from 1998 to 2006 in 47 large marine ecosystems around the world were found to be significantly related to: changes in estimated cholorophyll a (a proxy for phytoplankton biomass) in 18 of these ecosystems (mostly tropical and eastern boundary upwelling systems); changes in SST in 12 of these ecosystems (mostly mid-latitude); and changes in fishing intensity in 16 of these ecosystems (widely spread) (McOwen et al., 2015). Analysis of population data since the 1950s for 262 fish stocks across 39 large marine ecosystems and the high seas suggest that average recruitment to the stocks has declined by around 3% of the historical maximum per decade with variations between regions and stocks (Britten et al., 20161219). The declines (69% of the studied stocks, 31 of the 39 assessed large marine ecosystems) are significantly related to estimated chlorophyll a concentration and the intensity of fishing, with the North Atlantic showing the steepest declines (Britten et al., 20161220). In addition, recent meta-analysis of population data from 235 fish stocks worldwide from 1930 to 2010 suggest that the maximum catch potential from these populations decreased by 4.1% (95% confidence span 9.0% decline to 0.3% increase) during this period with variations between fish stocks and regions (Free et al., 20191221). Specifically, temperature is a significant factor explaining changes in catch potential of 12% of the fish stocks, with East Asian regions having the largest stock declines related to warming. In intermediate latitudes across the Atlantic, Indian and Pacific Oceans, catches of tropical tunas, including skipjack and yellowfin tuna, are significantly and positively related to increases in SST, although the overall catches across latitudinal zones do not show significant change (Monllor-Hurtado et al., 20171222). Observational evidence from spatial and temporal linkages between catches and oceanographic variables therefore supports the conclusions from AR5 WGII and SR15 that potential fisheries catches have already been impacted by the effects of warming and changing primary production on growth, reproduction and survival of fish stocks (robust evidence, high agreement, high confidence).