Toward a Solution of the “Peruvian Puzzle”: Pelagic Food-Web Structure and Trophic Interactions in the Northern Humboldt Current Upwelling System Off Peru

The northern Humboldt Current upwelling system (HCS) belongs to the most productive marine ecosystems, providing five to eight times higher fisheries landings per unit area than other coastal upwelling systems. To solve this “Peruvian puzzle”, to elucidate the pelagic food-web structure and to better understand trophic interactions in the HCS, a combined stable isotope and fatty acid trophic biomarker approach was adopted for key zooplankton taxa and higher trophic positions with an extensive spatial coverage from 8.5 to 16°S and a vertical range down to 1,000 m depth. A pronounced regional shift by up to ∼5‰ in the δ15N baseline of the food web occurred from North to South. Besides regional shifts, δ15N ratios of particulate organic matter (POM) also tended to increase with depth, with differences of up to 3.8‰ between surface waters and the oxygen minimum zone. In consequence, suspension-feeding zooplankton permanently residing at depth had up to ∼6‰ higher δ15N signals than surface-living species or diel vertical migrants. The comprehensive data set covered over 20 zooplankton taxa and indicated that three crustacean species usually are key in the zooplankton community, i.e., the copepods Calanus chilensis at the surface and Eucalanus inermis in the pronounced OMZ and the krill Euphausia mucronata, resulting in an overall low number of major trophic pathways toward anchovies. In addition, the semi-pelagic squat lobster Pleuroncodes monodon appears to play a key role in the benthic-pelagic coupling, as indicated by highest δ13C’ ratios of −14.7‰. If feeding on benthic resources and by diel vertical migration, they provide a unique pathway for returning carbon and energy from the seafloor to the epipelagic layer, increasing the food supply for pelagic fish. Overall, these mechanisms result in a very efficient food chain, channeling energy toward higher trophic positions and partially explaining the “Peruvian puzzle” of enormous fish production in the HCS.

Marine heatwaves in the Humboldt current system: from 5-day localized warming to year-long El Niños

During the last 4 decades punctual occurrences of extreme ocean temperatures, known as marine heatwaves (MHWs), have been regularly disrupting the coastal ecosystem of the Peru-Chile eastern boundary upwelling system. In fact, this coastal system and biodiversity hot-spot is regularly impacted by El Niño events, whose variability has been related to the longest and most intense MHWs in the world ocean. However the intensively studied El Niños tend to overshadow the MHWs of shorter duration that are significantly more common in the region. Using sea surface temperature data from 1982 to 2019 we investigate the characteristics and evolution of MHWs, distinguishing events by duration. Results show that long duration MHWs (> 100 days) preferentially affect the coastal domain north of 15° S and have decreased in both occurrence and intensity in the last four decades. On the other hand, shorter events, which represent more than 90% of all the observed MHWs, are more common south of 15° S and show an increase in their thermal impact as well as on the number of affected days, particularly those spanning 30–100 days. We also show that long duration MHWs variability in the coastal domain is well correlated with the remote equatorial variability while the onset of short events (< 10 days) generally goes along with a relaxation of the local coastal wind.

Vertebrate Palaeoecology of the Pisco Formation (Miocene, Peru): Glimpses into the Ancient Humboldt Current Ecosystem

The northward-flowing Humboldt Current hosts perpetually high levels of productivity along the western coast of South America. Here, we aim to elucidate the deep-time history of this globally important ecosystem based on a detailed palaeoecological analysis of the exceptionally preserved middle–upper Miocene vertebrate assemblages of the Pisco Formation of the East Pisco Basin, southern Peru. We summarise observations on hundreds of fossil whales, dolphins, seals, seabirds, turtles, crocodiles, sharks, rays, and bony fishes to reconstruct ecological relationships in the wake of the Middle Miocene Climatic Optimum, and the marked cooling that followed it. The lowermost, middle Miocene Pisco sequence (P0) and its vertebrate assemblage testify to a warm, semi-enclosed, near-shore palaeoenvironment. During the first part of the Tortonian (P1), high productivity within a prominent upwelling system supported a diverse assemblage of mesopredators, at least some of which permanently resided in the Pisco embayment and used it as a nursery or breeding/calving area. Younger portions of the Pisco Formation (P2) reveal a more open setting, with wide-ranging species like rorquals increasingly dominating the vertebrate assemblage, but also local differences reflecting distance from the coast. Like today, these ancient precursors of the modern Humboldt Current Ecosystem were based on sardines, but notably differed from their present-day equivalent in being dominated by extremely large-bodied apex predators like Livyatan melvillei and Carcharocles megalodon.

Nitrogen loss processes in response to upwelling in a Peruvian coastal setting dominated by denitrification – a mesocosm approach

Upwelling of nutrient-rich deep waters make eastern boundary upwelling systems (EBUSs), such as the Humboldt Current system, hot spots of marine productivity. Associated settling of organic matter to depth and consecutive aerobic decomposition results in large subsurface water volumes being oxygen depleted. Under these circumstances, organic matter remineralisation can continue via denitrification, which represents a major loss pathway for bioavailable nitrogen. Additionally, anaerobic ammonium oxidation can remove significant amounts of nitrogen in these areas. Here we assess the interplay of suboxic water upwelling and nitrogen cycling in a manipulative offshore mesocosm experiment. Measured denitrification rates in incubations with water from the oxygen-depleted bottom layer of the mesocosms (via 15N label incubations) mostly ranged between 5.5 and 20 nmol N2 L−1 h−1 (interquartile range), reaching up to 80 nmol N2 L−1 h−1. However, actual in situ rates in the mesocosms, estimated via Michaelis–Menten kinetic scaling, did most likely not exceed 0.2–4.2 nmol N2 L−1 h−1 (interquartile range) due to substrate limitation. In the surrounding Pacific, measured denitrification rates were similar, although indications of substrate limitation were detected only once. In contrast, anammox (anaerobic ammonium oxidation) made only a minor contribution to the overall nitrogen loss when encountered in both the mesocosms and the Pacific Ocean. This was potentially related to organic matter C / N stoichiometry and/or process-specific oxygen and hydrogen sulfide sensitivities. Over the first 38 d of the experiment, total nitrogen loss calculated from in situ rates of denitrification and anammox was comparable to estimates from a full nitrogen budget in the mesocosms and ranged between ∼ 1 and 5.5 µmol N L−1. This represents up to ∼ 20 % of the initially bioavailable inorganic and organic nitrogen standing stocks. Interestingly, this loss is comparable to the total amount of particulate organic nitrogen that was exported into the sediment traps at the bottom of the mesocosms at about 20 m depth. Altogether, this suggests that a significant portion, if not the majority of nitrogen that could be exported to depth, is already lost, i.e. converted to N2 in a relatively shallow layer of the surface ocean, provided that there are oxygen-deficient conditions like those during coastal upwelling in our study. Published data for primary productivity and nitrogen loss in all EBUSs reinforce such conclusion.

Latitudinal changes in the lipid content and fatty acid profiles of juvenile female red squat lobsters (Pleuroncodes monodon) in breeding areas of the Humboldt Current System

The red squat lobster Pleuroncodes monodon is a species of high commercial value that inhabits the Humboldt Current System. Along the Chilean coast, two populations are exploited by the fishing industry, one located off the coast of Coquimbo and the other off the coast of Concepción. Yet, it is unknown whether there are differences in the “bioenergetic fuel” (measured as lipid content and fatty acid profile) of juvenile populations of these two fishing units and whether these bioenergetic compounds can be modulated by differences in the environmental parameters (such as temperature or chlorophyll-a) of their breeding areas. To shed some light on this, we measured the lipid content and fatty acid profiles of the viscera and muscle of juvenile female red squat lobsters from these two fishing units, specifically from breeding areas near long-exploited fishing grounds: a) the northern fishing unit (NFU, from 26°S to 30°S) and b) the southern fishing unit (SFU, from 32°S to 37°S). We found differences in the lipid content, fatty acid profiles, and ratios of saturated fatty acids (C16:0/C18:0) of juvenile females from these two locations. In addition, the essential fatty acids (DHA/EPA) found in the viscera versus the muscle of these lobsters varied significantly. Juvenile females from the SFU (i.e. Concepción) showed a higher lipid content compared to the juvenile females from the NFU (i.e. Coquimbo). Consistently, individuals from the SFU had a higher content of fatty acids, which also proved to be richer in saturated and monounsaturated fatty acids compared to those from the NFU. Our results are important for the fisheries in both areas because these juvenile populations are the source of new recruits for the adult populations that are exploited by the fishing industry. Our study also aids in determining which populations are healthier or of better quality in bioenergetic terms. Furthermore, increasing the incorporation of bioenergetic parameters in fishery models is essential for the recruitment and stock assessment within an ecosystem approach, since it allows for the evaluation of the nutritional condition of different fishing populations.