The vulnerability of the marine ecosystem to climate change is considered to be a function of exposure and sensitivity to stressors, combined with adaptive capacity53,54. In this study, we quantify one part of this equation—the potential for exposure to environmental drivers. Marine ecosystems, and the individual organisms that make up those ecosystems, are adapted to the range of conditions they experience33,34. When the environment changes sufficiently that new conditions, or a new combination of conditions, emerge and persist, the organisms must adapt, migrate to more favourable areas, or face extinction. In this way, climate forcing can induce changes at the organism level, which result in changes to ecosystem structure, species interactions and food web dynamics55,56. If novel conditions emerge rapidly, species have less time to either adapt or migrate, potentially increasing the probability that disruption to the ecosystem will occur.

We find that the climate change-driven trend in pH already exceeds the range in natural seasonal variability over most of the ocean, as does SST in the subtropics and Arctic. For PP and interior oxygen content, although the trend is large, the natural variability is also large, resulting in later ToE. Species adapted to living in regions of low variability are likely to have relatively narrow environmental niches8 and may be living close to their maximum tolerance57. Subtropical and tropical species are therefore likely to be more sensitive to the rapid emergence of climate change trends. Polar species are also particularly vulnerable as they cannot shift their geographical range northward in response to emerging drivers and so must either adapt to changing conditions or go extinct58. However, new ecological niches may open for species resilient to ocean acidification and warming waters and inured to the large natural variability that occurs in PP and oxygen.

Our model results suggest that seasonal minimum pH levels have been lower than the previously experienced natural range for >90 years already. This is consistent with analyses of (shorter) observational records, which suggest that trends in ocean acidification are likely to be anthropogenically driven50,59. However, evidence that this change has had a significant or lasting impact on marine organisms is scarce, except perhaps for warm-water coral communities60. Does this imply that the marine ecosystem is actually rather resilient to climate change? The key factor may be the speed with which climate change emerges in marine ecosystem drivers relative to the speed with which organisms can adapt. Individual species can seemingly adapt relatively rapidly (compared with the timescales of climate change) to new conditions. For example, a tropical reef fish was found to acclimate to acute exposure to warmer temperatures within two generations (damselfish have lifespans of >5 years61). Organisms with shorter lifespans, such as phytoplankton (∼few days), adapt correspondingly more rapidly. Coccolithophores, for example, adapted to a large degree to more acidic conditions within 500 generations62. The fossil record can also provide some insight into the possible impacts of climate change on marine ecosystems (although the paleoclimate changed at slower rates than projected for anthropogenic climate change). During the Paleocene–Eocene Thermal Maximum for example, up to 50% of benthic foraminifera went extinct63 and warm-water species expanded their ranges northward64,65. Although the limits of adaptation capacity are presently very poorly known, past extinctions at slower rates of climate change suggest that adaptation rates in some organisms are unlikely to be fast enough to keep pace, ultimately implying extinction.

We demonstrate that a heterogeneous mosaic of multiple environmental drivers develops in the next 50 years (Fig. 2c). This mosaic suggests that species resilient to change in one driver but negatively affected by another may be able to migrate to newly formed suitable habitats, provided the velocity of climate change (sensu16) does not outpace migration speed. In addition, some species may be able to alter their depth range so that they avoid decreases in thermocline oxygen whilst still remaining within their thermal niche (note that here we only assess changes in surface temperature). Again, little is known about the potential migration speed of marine organisms. Large motile species such as fishes and mammals are likely to be able to migrate rapidly to more favourable conditions (although locating these refugia may not be as simple as tracking a northward-moving isotherm). Smaller motile species, such as zooplankton, have also been observed to migrate in response to climate trends or variability, as in the North Atlantic where the distribution of warm-water copepod species has shifted northward in recent decades66. Planktonic species may be able to rely on rapid dispersal to maintain populations, whereas sessile species may not be able to migrate sufficiently rapidly to keep pace with future climate change. Returning to the Paleocene–Eocene Thermal Maximum as an analogy, mobile crustaceans avoided significant community changes67, whereas sedentary sediment dwellers were heavily impacted63.

Translating the emergence of climate drivers that we present here into an understanding of how the structure and functioning of the marine ecosystem may respond is an extremely challenging task. The interplay between biogeochemical stressors, including synergistic effects, adaptation and migration potential and speed, the bioclimatic envelope, organisms’ climate sensitivity, non-linear responses to changing conditions, short-term acclimatization and extinction risk is complex in the extreme. Even the first step, of connecting the emergence of climate change in ocean conditions with potential stress in marine organisms, requires the implicit assumption that the niche width of an organism scales with the local variability it experiences, and if conditions exceed that variability then a response (whether negative or positive) may occur. However, many species appear to thrive in environments that are less than optimal, for example, some marine fish and invertebrates have warmer or cooler temperature optima than the environment in which they are found54. In addition, the emergence of persistent, anomalously low oxygen (compared to previous seasonal variability) may not be of relevance to individual organisms if concentrations still remain above hypoxic levels, although oxygen has been found to limit animal life even at higher concentrations8.

Currently, it is not clear whether ecosystem-wide adaptation or migration can outpace the speed at which multiple drivers emerge (Fig. 1). How individual species will fare, or how the ecosystem as the sum of its parts will fare, is poorly understood. What is clear however is that there are likely to be winners and losers in the future ocean68. However, lacking the ability to predict the future impact of drivers on marine ecosystems creates significant challenges to determining an appropriate course for sustainable management of the ecosystem services, such as fisheries, that they provide. The quantification of the ToE of multiple drivers presented here is an important first step in achieving an understanding of the response of marine ecosystems to future climate change.