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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The Humboldt Current System is the most productive upwelling system globally in terms of fish and sustains huge populations of guano-producing birds as well as the world's largest single-species fishery targeting the Peruvian anchovy Engraulis ringens. Peruvian boobies Sula variegata and Guanay cormorants Phalacrocorax bougainvillii are the most abundant seabird species in the system and feed mainly on anchovies. By using high precision GPS and time−depth recorders, we compared the horizontal and vertical movements of the 2 species to examine whether segregation occurs between their foraging areas and whether there is any evidence for prey depletion close to large colonies. A simultaneous acoustic cruise in the foraging area of the 2 bird species estimated the vertical and horizontal distributions of anchovy schools. During the study period in austral spring 2008, the 2 bird species foraged in the same area, at relatively short distance from the colony (average, 19 to 20 km). Both species foraged at similar times of the day, although Peruvian boobies tended to leave earlier in the morning and return later in the evening than did the cormorants. Foraging trips were similar in duration, distance covered and range. Peruvian anchovies were abundant and schools occurred at shallow depths (median, 7 m; range, 3 to 34 m), mainly to the north and west of the colony where foraging bouts of the 2 species overlapped extensively. Cormorants are mainly pelagic surface divers, diving at shallow depths (median, 4 m) but reaching up to 32 m depth, and are thus able to track all the depths at which anchovies occurred during the study period. Peruvian boobies are plunge divers able to reach occasionally 10 m (median, 2 m), and are thus able to reach only shallow anchovy schools. No sexspecific differences in horizontal or vertical movements occurred between males and females in the 2 species. We suggest that the absence of significant differences between horizontal movements of boobies and cormorants, the relative short duration of their trips and the high number of chicks fledged were probably explained by the high abundance and accessibility of anchovies. Differences may be exacerbated during years of low anchovy abundance or accessibility, especially El Niño years. KEY WORDS: Peruvian anchovy · Telemetry · Phalacrocorax bougainvillii · Sula variegataResale or republication not permitted without written consent of the publisher

ABSTRACT. Changes in abundance and distribution of anchovy and other species of pelagic fish of the Northern Humboldt Current System (NHCS) are driven by environmental forcing variations in different time and spatial scales between the coastal 'cold' ecosystem and the 'warm' oceanic one. Data to study these fluctuations have come mostly from the fishery to show how anchovy (Engraulis ringens) increases when sardine (Sardinops sagax) declines and vice versa. However, using acoustic data on latitudinal biomass we show that other species as mackerel (Scomber japonicus) and jack mackerel (Trachurus murphyi) also follow the same opposed trend, then the fishery data hides the true dimension of the balance of abundance among species. Based on Hovmoller diagrams we scrutinized the changes in interannual latitudinal acoustic biomass, landings and influence of El Niño events from 1966 to 2009 in order to describe: 1) how the anchovy decadal distribution pattern moved from south to north since the 1960's; 2) how there have been produced concomitant changes in the latitudinal abundance and distribution of other species such as sardine, jack mackerel and mackerel before, during and after El Niño events; and 3) what was the overall effect of the succession of El Niño events on all these pelagic species. We concluded that: a) every El Niño event has had an effect on the expansion or contraction of pelagic species distribution and abundance, with different latitudinal effects; and b) the El Niño 1997-98 did not trigger but accelerated a decline phase on the abundance of sardine, jack mackerel and mackerel by a reduction of their ideal habitat due to an expansion of the coastal ecosystem caused by a shallower location of the upper limit of the Oxygen Minimum Zone (OMZ) These findings observed using past data might be taken into consideration for fishery management purposes when considering future scenarios. Keywords: acoustics, small pelagic fish, landings, biomass, latitudinal distribution, El Niño, Perú. Tendencias espacio-temporales en la distribución de la biomasa de anchoveta peruana y de otros peces pelágicos pequeños entre 1966 y 2009RESUMEN. Los cambios en la abundancia y distribución de anchoveta y de otras especies de pequeños peces pelágicos de la región norte del Sistema de la Corriente de Humboldt (NHCS) son el producto de la variación de forzantes ambientales en diferentes escalas de tiempo y espacio que influyen entre el ecosistema 'frío' costero y el oceánico 'cálido'. La información para estudiar estas fluctuaciones provienen mayormente de las pesquerías, y muestran que la anchoveta (Engraulis ringens) incrementa su abundancia cuando la de sardina (Sardinops sagax) declina y viceversa. Sin embargo, utilizando datos acústicos de biomasa latitudinal se muestra que otras especies como la caballa (Scomber japonicus) y el jurel (Trachurus murphyi) también siguen la misma tendencia opuesta a anchoveta, lo que en principio indica que los datos pesqueros disponibles no indican la verdadera dimensión del balance de abund...

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