Spatial variation in environmental conditions can elicit predictable size and morphological responses in marine organisms through influences on physiology. Thus, spatial and temporal variation in marine organism size and shape are often used to infer paleoenvironmental conditions, such as dissolved oxygen concentrations. Benthic foraminifera commonly serve as a tool for reconstructing past ocean oxygen levels. For example, benthic foraminiferal species assemblages, within- and among-species patterns of test morphology, and geochemical analyses of carbonate tests are often employed to reconstruct past marine conditions. In this study, we measured the sizes and shapes of modern foraminifera representing four species that inhabit a steep oxygen gradient in the Santa Monica Basin on the Southern California Borderland with the aim of quantifying the influence of oxygen availability on foraminiferal morphology, both within and among species. Most foraminifera rely on aerobic respiration, but the four benthic foraminifera from the Santa Monica Basin do not show the predicted size and morphological responses to variations in dissolved oxygen concentrations based on first principles of cell physiology: Bolivina spissa shows no volume or volume-to-surface area response, Uvigerina peregrina increases in both test volume and volume-to-surface area ratio with decreasing dissolved oxygen concentrations, and both Bolivina argentea and Loxostomum pseudobeyrichi decrease in test volume, but only L.pseudobeyrichi shows a decrease in test volume-to-surface area ratio with decreasing oxygen concentrations as expected from physiological predictions. These findings imply that the morphological responses of individual foraminiferal species are not necessarily representative of the responses of other foraminiferal species within the community. Our findings further suggest that these species use physiological strategies such as depressed metabolic rates and alternative energy metabolisms to persist in low oxygen environments and, therefore, cannot be used in any simple way as paleo-oxygen indicators. Should Proterozoic (1,000-542 million years ago) protists have possessed metabolic strategies similar to foraminifera, the sizes and shapes of protists in the fossil record may not usefully constrain ambient oxygen conditions during the appearance and initial taxonomic radiation of heterotrophic eukaryotes.
