El Niño and similar perturbation effects on the benthos of the Humboldt, California, and Benguela Current upwelling ecosystems

To a certain degree, Eastern Boundary Current (EBC) ecosystems are similar: Cold bottom water from moderate depths, rich in nutrients, is transported to the euphotic zone by a combination of trade winds, Coriolis force and Ekman transport. The resultant high primary production fuels a rich secondary production in the upper pelagic and nearshore zones, but where O2 exchange is restricted, it creates oxygen minimum zones (OMZs) at shelf and upper slope (Humboldt and Benguela Current) or slope depths (California Current). These hypoxic zones host a specifically adapted, small macroand meiofauna together with giant sulphur bacteria that use nitrate to oxydise H2S. In all EBC, small polychaetes, large nematodes and other opportunistic benthic species have adapted to the hypoxic conditions and co-exist with sulphur bacteria, which seem to be particularly dominant off Peru and Chile. However, a massive reduction of macrobenthos occurs in the core of the OMZ. In the Humboldt Current area the OMZ ranges between <100 and about 600 m, with decreasing thickness in a poleward direction. The OMZ merges into better oxygenated zones towards the deep sea, where large cold-water megaand macrofauna occupy a dominant role as in the nearshore strip. The Benguela Current OMZ has a similar upper limit but remains shallower. It also hosts giant sulphur bacteria but little is known about the benthic fauna. However, sulphur eruptions and intense hypoxia might preclude the coexistence of significant megaund macrobenthos. Conversely, off North America the upper limit of the OMZ is considerably deeper (e.g., 500– Correspondence to: W. E. Arntz ([email protected]) 600 m off California and Oregon), and the lower boundary may exceed 1000 m. The properties described are valid for very cold and cold (La Niña and “normal”) ENSO conditions with effective upwelling of nutrient-rich bottom water. During warm (El Niño) episodes, warm water masses of low oxygen concentration from oceanic and equatorial regions enter the upwelling zones, bringing a variety of (sub)tropical immigrants. The autochthonous benthic fauna emigrates to deeper water or poleward, or suffers mortality. However, some local macrofaunal species experience important population proliferations, presumably due to improved oxygenation (in the southern hemisphere), higher temperature tolerance, reduced competition or the capability to use different food. Both these negative and positive effects of El Niño influence local artisanal fisheries and the livelihood of coastal populations. In the Humboldt Current system the hypoxic seafloor at outer shelf depths receives important flushing from the equatorial zone, causing havoc on the sulphur bacteria mats and immediate recolonisation of the sediments by megaand macrofauna. Conversely, off California, the intruding equatorial water masses appear to have lower oxygen than ambient waters, and may cause oxygen deficiency at upper slope depths. Effects of this change have not been studied in detail, although shrimp and other taxa appear to alter their distribution on the continental margin. Other properties and reactions of the two Pacific EBC benthic ecosystems to El Niño seem to differ, too, as does the overall impact of major episodes (e.g., 1982/1983(1984) vs. 1997/1998). The relation of the “Benguela Niño” to ENSO seems unclear although many 244 W. E. Arntz et al.: El Niño and similar perturbation effects Pacific-Atlantic ocean and atmosphere teleconnections have been described. Warm, low-oxygen equatorial water seems to be transported into the upwelling area by similar mechanisms as in the Pacific, but most major impacts on the eukaryotic biota obviously come from other, independent perturbations such as an extreme eutrophication of the sediments ensuing in sulphidic eruptions and toxic algal blooms. Similarities and differences of the Humboldt and California Current benthic ecosystems are discussed with particular reference to ENSO impacts since 1972/73. Where there are data available, the authors include the Benguela Current ecosystem as another important, non-Pacific EBC, which also suffers from the effects of hypoxia.