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Abstract The Southern Ocean is a region of high surface nutrient content, reflecting an inefficient biological carbon pump. The variability, predictability, and causes of changes in these nutrient levels on interannual to decadal time scales remain unclear. We employ a deep learning approach, specifically a Temporal Convolution Attention Neural Network (TCANN), to conduct multi‐year forecasting of surface based on oceanic physical drivers. The TCANN successfully replicates testing data with a prediction skill extending to at least 4 years with the GFDL‐ESM4‐driven model and 1 year with the observation‐driven model. To benchmark the results, we compare the prediction skill of TCANN with a simple persistence model and two regression methods, a linear regression and a ridge regression. The TCANN model was able to predict variability with a higher skill than persistence and the two regression methods indicating that non‐linearities present in the system become too high to predict inter‐annual variability with traditional regression methods. To enhance the interpretability of the predictions, we explore three explainable AI techniques: occlusion analysis, integrated gradients, and Gradient Shap. The outcomes suggest a crucial role played by salinity processes and buoyancy/potential density fluxes on the prediction of on annual time scales. The deep learning tools' ability to provide skillful forecasts well into the future presents a promising avenue for gaining insights into how the Southern Ocean's surface nutrients respond to climate change based on physical quantities. 

Fossil assemblages exhibit a global depletion in taxonomic distinctiveness in the aftermath of the end-Permian mass extinction (~252 million years ago), but little is known about why. Here, we examine whether biotic homogenization can be explained by tropical survivors tracking an expansion of their preferred habitat, measured in terms of the ratio of environmental oxygen supply to metabolic demand. We compare spatial similarity in community composition among marine invertebrate fossils represented by bivalve and gastropod fossils with predictions from an ecophysiological model of habitat that diagnoses areas in the ocean that can sustain the aerobic requirements of marine invertebrates. Modeled biogeographic responses to climate change yield an increase in global similarity of community composition among surviving ecophysiotypes, consistent with patterns in the fossil record and arguing for a physiological control on earliest Triassic biogeography. 

The minimum O2 needed to fuel the demand of aquatic animals is commonly observed to increase with temperature, driven by accelerating metabolism. However, recent measurements of critical O2 thresholds (“Pcrit”) reveal more complex patterns, including those with a minimum at an intermediate thermal “optimum”. To discern the prevalence, physiological drivers, and biogeographic manifestations of such curves, we analyze new experimental and biogeographic data using a general dynamic model of aquatic water breathers. The model simulates the transfer of oxygen from ambient water through a boundary layer and into animal tissues driven by temperature-dependent rates of metabolism, diffusive gas exchange, and ventilatory and circulatory systems with O2-protein binding. We find that a thermal optimum in Pcrit can arise even when all physiological rates increase steadily with temperature. This occurs when O2 supply at low temperatures is limited by a process that is more temperature sensitive than metabolism, but becomes limited by a less sensitive process at warmer temperatures. Analysis of published species respiratory traits suggests that this scenario is not uncommon in marine biota, with ventilation and circulation limiting supply under cold conditions and diffusion limiting supply at high temperatures. Using occurrence data, we show that species with these physiological traits inhabit lowest O2 waters near the optimal temperature for hypoxia tolerance and are restricted to higher O2 at temperatures above and below this optimum. Our results imply that hypoxia tolerance can decline under both cold and warm conditions and thus may influence both poleward and equatorward species range limits. 

Abstract In an ocean that is rapidly warming and losing oxygen, accurate forecasting of species’ responses must consider how this environmental change affects fundamental aspects of their physiology. Here, we develop an absolute metabolic index (Φ A ) that quantifies how ocean temperature, dissolved oxygen and organismal mass interact to constrain the total oxygen budget an organism can use to fuel sustainable levels of aerobic metabolism. We calibrate species-specific parameters of Φ A with physiological measurements for red abalone ( Haliotis rufescens ) and purple urchin ( Strongylocentrotus purpuratus ). Φ A models highlight that the temperature where oxygen supply is greatest shifts cooler when water loses oxygen or organisms grow larger, providing a mechanistic explanation for observed thermal preference patterns. Viable habitat forecasts are disproportionally deleterious for red abalone, revealing how species-specific physiologies modulate the intensity of a common climate signal, captured in the newly developed Φ A framework. 

The global ocean's oxygen content has declined significantly over the past several decades and is expected to continue decreasing under global warming, with far-reaching impacts on marine ecosystems and biogeochemical cycling. Determining the oxygen trend, its spatial pattern, and uncertainties from observations is fundamental to our understanding of the changing ocean environment. This study uses a suite of CMIP6 Earth system models to evaluate the biases and uncertainties in oxygen distribution and trends due to sampling sparseness. Model outputs are sub-sampled according to the spatial and temporal distribution of the historical shipboard measurements, and the data gaps are filled by a simple optimal interpolation method using Gaussian covariance with a constant e-folding length scale. Sub-sampled results are compared to full model output, revealing the biases in global and basin-wise oxygen content trends. The simple optimal interpolation underestimates the modeled global deoxygenation trends, capturing approximately two-thirds of the full model trends. The North Atlantic and subpolar North Pacific are relatively well sampled, and the simple optimal interpolation is capable of reconstructing more than 80% of the oxygen trend in the non-eddying CMIP models. In contrast, pronounced biases are found in the equatorial oceans and the Southern Ocean, where the sampling density is relatively low. The application of the simple optimal interpolation method to the historical dataset estimated the global oxygen loss to be 1.5% over the past 50 years. However, the ratio of the global oxygen trend between the sub-sampled and full model output has increased the estimated loss rate in the range of 1.7% to 3.1% over the past 50 years, which partially overlaps with previous studies. The approach taken in this study can provide a framework for the intercomparison of different statistical gap-filling methods to estimate oxygen content trends and their uncertainties due to sampling sparseness. 

Abstract The proportion of major elements in marine organic matter links cellular processes to global nutrient, oxygen and carbon cycles. Differences in the C:N:P ratios of organic matter have been observed between ocean biomes, but these patterns have yet to be quantified from the underlying small-scale physiological and ecological processes. Here we use an ecosystem model that includes adaptive resource allocation within and between ecologically distinct plankton size classes to attribute the causes of global patterns in the C:N:P ratios. We find that patterns of N:C variation are largely driven by common physiological adjustment strategies across all phytoplankton, while patterns of N:P are driven by ecological selection for taxonomic groups with different phosphorus storage capacities. Although N:C varies widely due to cellular adjustment to light and nutrients, its latitudinal gradient is modest because of depth-dependent trade-offs between nutrient and light availability. Strong latitudinal variation in N:P reflects an ecological balance favouring small plankton with lower P storage capacity in the subtropics, and larger eukaryotes with a higher cellular P storage capacity in nutrient-rich high latitudes. A weaker N:P difference between southern and northern hemispheres, and between the Atlantic and Pacific oceans, reflects differences in phosphate available for cellular storage. Despite simulating only two phytoplankton size classes, the emergent global variability of elemental ratios resembles that of all measured species, suggesting that the range of growth conditions and ecological selection sustain the observed diversity of stoichiometry among phytoplankton. 

Abstract A central question in the study of mass extinction is whether these events simply intensify background extinction processes and patterns versus change the driving mechanisms and associated patterns of selectivity. Over the past two decades, aided by the development of new fossil occurrence databases, selectivity patterns associated with mass extinction have become increasingly well quantified and their differences from background patterns established. In general, differences in geographic range matter less during mass extinction than during background intervals, while differences in respiratory and circulatory anatomy that may correlate with tolerance to rapid change in oxygen availability, temperature, and pH show greater evidence of selectivity during mass extinction. The recent expansion of physiological experiments on living representatives of diverse clades and the development of simple, quantitative theories linking temperature and oxygen availability to the extent of viable habitat in the oceans have enabled the use of Earth system models to link geochemical proxy constraints on environmental change with quantitative predictions of the amount and biogeography of habitat loss. Early indications are that the interaction between physiological traits and environmental change can explain substantial proportions of observed extinction selectivity for at least some mass extinction events. A remaining challenge is quantifying the effects of primary extinction resulting from the limits of physiological tolerance versus secondary extinction resulting from the loss of taxa on which a given species depended ecologically. The calibration of physiology-based models to past extinction events will enhance their value in prediction and mitigation efforts related to the current biodiversity crisis. 

Rising temperatures are associated with reduced body size in many marine species, but the biological cause and generality of the phenomenon is debated. We derive a predictive model for body size responses to temperature and oxygen (O 2 ) changes based on thermal and geometric constraints on organismal O 2 supply and demand across the size spectrum. The model reproduces three key aspects of the observed patterns of intergenerational size reductions measured in laboratory warming experiments of diverse aquatic ectotherms (i.e., the “temperature-size rule” [TSR]). First, the interspecific mean and variability of the TSR is predicted from species’ temperature sensitivities of hypoxia tolerance, whose nonlinearity with temperature also explains the second TSR pattern—its amplification as temperatures rise. Third, as body size increases across the tree of life, the impact of growth on O 2 demand declines while its benefit to O 2 supply rises, decreasing the size dependence of hypoxia tolerance and requiring larger animals to contract by a larger fraction to compensate for a thermally driven rise in metabolism. Together our results support O 2 limitation as the mechanism underlying the TSR, and they provide a physiological basis for projecting ectotherm body size responses to climate change from microbes to macrofauna. For small species unable to rapidly migrate or evolve greater hypoxia tolerance, ocean warming and O 2 loss in this century are projected to induce >20% reductions in body mass. Size reductions at higher trophic levels could be even stronger and more variable, compounding the direct impact of human harvesting on size-structured ocean food webs. 

ABSTRACT Tropical reef ecosystems are strongly influenced by the composition of coral species, but the factors influencing coral diversity and distributions are not fully understood. Here we demonstrate that large variations in the relative abundance of three major coral species across adjacent Caribbean reef sites are strongly related to their different low O₂tolerances. In laboratory experiments designed to mimic reef conditions, the cumulative effect of repeated nightly low O₂drove coral bleaching and mortality, with limited modulation by temperature. After four nights of repeated low O₂, species responses also varied widely, from > 50% bleaching inAcropora cervicornisto no discernable sensitivity ofPorites furcata.A simple metric of hypoxic pressure that combines these experimentally derived species sensitivities with high‐resolution field data accurately predicts the observed relative abundance of species across three reefs. Only the well‐oxygenated reef supported the framework‐building hypoxia‐sensitiveAcropora cervicornis, while the hypoxia‐tolerant weedy speciesPorites furcatawas dominant on the most frequently O₂‐deplete reef. Physiological exclusion of acroporids from these O₂‐deplete reefs underscores the need for hypoxia management to reduce extirpation risk.