BackgroundIn the southeastern tropical Pacific anchovy (Engraulis ringens) and sardine (Sardinops sagax) abundance have recently fluctuated on multidecadal scales and food and temperature have been proposed as the key parameters explaining these changes. However, ecological and paleoecological studies, and the fact that anchovies and sardines are favored differently in other regions, raise questions about the role of temperature. Here we investigate the role of oxygen in structuring fish populations in the Peruvian upwelling ecosystem that has evolved over anoxic conditions and is one of the world's most productive ecosystems in terms of forage fish. This study is particularly relevant given that the distribution of oxygen in the ocean is changing with uncertain consequences.Methodology/Principal FindingsA comprehensive data set is used to show how oxygen concentration and oxycline depth affect the abundance and distribution of pelagic fish. We show that the effects of oxygen on anchovy and sardine are opposite. Anchovy flourishes under relatively low oxygen conditions while sardine avoid periods/areas with low oxygen concentration and restricted habitat. Oxygen consumption, trophic structure and habitat compression play a fundamental role in fish dynamics in this important ecosystem.Conclusions/SignificanceFor the ocean off Peru we suggest that a key process, the need to breathe, has been neglected previously. Inclusion of this missing piece allows the development of a comprehensive conceptual model of pelagic fish populations and change in an ocean ecosystem impacted by low oxygen. Should current trends in oxygen in the ocean continue similar effects may be evident in other coastal upwelling ecosystems.

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Abstract. Over the last decades, the Humboldt Current upwelling ecosystem, particularly the northern component off the coast of Peru, has drawn the interest of the scientific community because of its unique characteristics: it is the upwelling system with the biggest catch productivity despite the fact it is embedded in a shallow and intense oxygen minimum zone (OMZ). It is also an area of intense nitrogen loss and anammox activity and experiences large interannual variability associated with the equatorial remote forcing. In this context, we examined the oceanographic and biogeochemical variability associated with the OMZ off central Peru from a monthly time series (1996)(1997)(1998)(1999)(2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009)(2010)(2011) recorded off the coast of Callao (12• 02 S, 77 • 29 W). The data reveal a rich spectrum of variability in the OMZ that includes frequencies ranging from seasonal to interannual scales. Due to the efficient oceanic teleconnection off Peru, the observed variability is interpreted in the light of an estimate of the equatorial Kelvin wave contribution to sea level anomalies considering the peculiarities of its vertical structure (i.e., the first two baroclinic modes). The span of the data set allows us to contrast two OMZ regimes. The "strong" regime is associated with the strong 1997-1998 equatorial Pacific El Niño, during which the OMZ adjusted to Kelvin-wave-induced downwelling conditions that switched off the upwelling and drastically reduced nutrient availability. The "weak" regime corresponds to the post-2000 period associated with the occurrence of moderate central Pacific El Niño events and enhanced equatorial Kelvin wave activity, in which mean upwelling conditions are maintained. It is shown that the characteristics of the coupling between physics and biogeochemistry is distinct between the two regimes with the weak regime being associated with a larger explained variance in biogeochemical properties not linearly related to the ENSO oceanic teleconnection. The data also reveal a long-term trend from 1999 corresponding to a deepening of the oxygendeficient waters and warming. The implications of our results for understanding the OMZ dynamics off Peru are discussed.