Ocean deoxygenation and copepods: Coping with oxygen minimum zone variability

Abstract

Increasing deoxygenation (loss of oxygen) of the ocean, including expansion of oxygen minimum zones (OMZs), is a potentially important consequence of global warming. We examined present-day variability of vertical distributions of 23 calanoid copepod species in the Eastern Tropical North Pacific (ETNP) living in locations with different water column oxygen profiles and OMZ intensity (lowest oxygen concentration and its vertical extent in a profile). Copepods and hydrographic data were collected in vertically stratified day and night MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System) tows (0-1000 m) during four cruises over a decade (2007-2017) that sampled four ETNP locations: Costa Rica Dome, Tehuantepec Bowl, and two oceanic sites further north (21-22 N) off Mexico. The sites had different vertical oxygen profiles: some with a shallow mixed layer, abrupt thermocline, and extensive very low oxygen OMZ core; and others with a more gradual vertical development of the OMZ (broad mixed layer and upper oxycline zone) and a less extensive OMZ core where oxygen was not as low. Calanoid copepod species (including examples from the genera Eucalanus, Pleuromamma, and Lucicutia) demonstrated different distributional strategies (implying different physiological characteristics) associated with this variability. We identified sets of species that (1) changed their vertical distributions and depth of maximum abundance associated with the depth and intensity of the OMZ and its oxycline inflection points; (2) shifted their depth of diapause; (3) adjusted their diel vertical migration, especially the nighttime upper depth; or (4) expanded or contracted their depth range within the mixed layer and upper part of the thermocline in association with the thickness of the aerobic epipelagic zone (habitat compression concept). These distribution depths changed by tens to hundreds of meters depending on the species, oxygen profile, and phenomenon. For example, at the lower oxycline, the depth of maximum abundance for Lucicutia hulsemannae shifted from &tild;1/4600 to &tild;1/4800 m, and the depth of diapause for Eucalanus inermis shifted from &tild;1/4500 to &tild;1/4775 m, in an expanded OMZ compared to a thinner OMZ, but remained at similar low oxygen levels in both situations. These species or life stages are examples of "hypoxiphilic" taxa. For the migrating copepod Pleuromamma abdominalis, its nighttime depth was shallow ( &tild;1/420 m) when the aerobic mixed layer was thin and the low-oxygen OMZ broad, but it was much deeper ( &tild;1/4100 m) when the mixed layer and higher oxygen extended deeper; daytime depth in both situations was &tild;1/4300 m. Because temperature decreased with depth, these distributional depth shifts had metabolic implications. The upper ocean to mesopelagic depth range encompasses a complex interwoven ecosystem characterized by intricate relationships among its inhabitants and their environment. It is a critically important zone for oceanic biogeochemical and export processes and hosts key food web components for commercial fisheries. Among the zooplankton, there will likely be winners and losers with increasing ocean deoxygenation as species cope with environmental change. Changes in individual copepod species abundances, vertical distributions, and life history strategies may create potential perturbations to these intricate food webs and processes. Present-day variability provides a window into future scenarios and potential effects of deoxygenation.