MARTIN THIEL ET AL.40% <strong>of</strong> <strong>the</strong> annual l<strong>and</strong>ings <strong>of</strong> <strong>the</strong> HCS (Escribano et al. 2003 <strong>and</strong> references <strong>the</strong>rein). Thisproduction also constitutes an important way <strong>of</strong> sequestering CO 2 <strong>and</strong> supports a high rate <strong>of</strong>particulate organic matter (POM) exported to depth (González et al. 2000a, Pantoja et al. 2004).This material, which is partly remineralised in <strong>the</strong> water column, streng<strong>the</strong>ns <strong>the</strong> oxygen-minimumzone (OMZ) <strong>and</strong> promotes biogeochemical anaerobic processes. In this sense, a sequence <strong>of</strong>mechanisms that are determined by <strong>the</strong> oceanographic conditions is regulating <strong>the</strong> chemistry <strong>of</strong> <strong>the</strong>water column <strong>and</strong> <strong>the</strong> sea bottom. These processes gain relevance in several aspects <strong>of</strong> materialexchange (i.e., gas fluxes as CO 2 <strong>and</strong> N 2 O), implying that this area could play a key role in <strong>the</strong>main global cycles (i.e., oceanic productivity, global warming, authigenic carbonatic <strong>and</strong> phosphoritemineral formation, etc.).Oxygen distribution <strong>and</strong> relevance in organic carbon remineralisationMost research efforts have focused on specific areas where high productivity cells are recognised,but <strong>the</strong> interactions with biogeochemical processes are still poorly understood. Along <strong>the</strong> nor<strong>the</strong>rnmargin <strong>of</strong> Chile, a well-developed OMZ is observed between 100 <strong>and</strong> 500 m water depth (Blancoet al. 2001). This zone is basically a mid-water feature since <strong>the</strong> topography <strong>of</strong> <strong>the</strong> margin precludes<strong>the</strong> OMZ to impinge on a large area <strong>of</strong> <strong>the</strong> bottom, except <strong>of</strong>f Ant<strong>of</strong>agasta (Mejillones) where ittouches <strong>the</strong> bottom from 50 m to 300 m depth. Normally <strong>the</strong> shelf is extremely narrow fromsou<strong>the</strong>rn Peru to nor<strong>the</strong>rn Chile (10–15 km) in comparison with <strong>central</strong> Peru <strong>and</strong> sou<strong>the</strong>rn-<strong>central</strong>Chile (40–60 km; Strub et al. 1998), where <strong>the</strong> OMZ extends over a wide area <strong>of</strong> <strong>the</strong> shelf, promotingdistinct biogeochemical processes (Gutiérrez 2000, Neira et al. 2001). This characteristic <strong>of</strong> nor<strong>the</strong>rnareas could thus affect pathways (i.e., aerobic or anaerobic) associated with organic matter (OM)degradation in <strong>the</strong> sediments, which is an important source <strong>of</strong> regenerated nutrients to <strong>the</strong> watercolumn. Off Mejillones, a high percentage (86%) <strong>of</strong> photosyn<strong>the</strong>tically produced particulatedprotein is being degraded within <strong>the</strong> upper 30 m <strong>of</strong> <strong>the</strong> water column (Pantoja et al. 2004), coincidingwith oxygenated waters. In consequence, OM reaching <strong>the</strong> sediments at greater depths is depleted<strong>of</strong> proteins. Over <strong>the</strong> shallower shelf sediments, where also preserved fish debris <strong>and</strong> bones arefound (Milessi et al. 2005), high pigment concentrations have been reported (42–100 µg g −1 <strong>of</strong>chloroplastic pigment equivalents in surface sediments; P. Muñoz et al. 2004a, 2005). Thus, <strong>the</strong>reis a narrow b<strong>and</strong> <strong>of</strong> inshore sediments that are enriched in fresh OM coming from <strong>the</strong> water column.The remineralisation <strong>of</strong> this material could generate an important flux <strong>of</strong> nutrients contributing t<strong>of</strong>ertilisation <strong>of</strong> <strong>the</strong> water column. Similar predictions can be made for o<strong>the</strong>r areas <strong>of</strong> high PP along<strong>the</strong> coast <strong>of</strong> nor<strong>the</strong>rn Chile, where upwelled waters containing preformed nutrients are enrichedwith recycled nutrients derived from <strong>the</strong> OM degradation in shelf sediments. The relevance <strong>of</strong> <strong>the</strong>sea floor in water column fertilisation <strong>and</strong> biological productivity as well as its relevance in <strong>the</strong>global carbon cycle has not been well examined. Some information is available for <strong>the</strong> role <strong>of</strong>sediments near <strong>the</strong> main upwelling centres, but nothing is known about <strong>the</strong> biogeochemical processesalong <strong>the</strong> large extent <strong>of</strong> <strong>the</strong> margin between <strong>the</strong> main upwelling centres mentioned here.The high OM degradation rates result in CO 2 supersaturated waters that, in combination with<strong>the</strong> CO 2 input from upwelled waters, favour <strong>the</strong> CO 2 flux from <strong>the</strong> ocean to <strong>the</strong> atmosphere. Somestudies <strong>of</strong>f Ant<strong>of</strong>agasta (23–24°S) <strong>and</strong> Coquimbo (30°S) have measured a saturation <strong>of</strong> >200% inupwelled waters, an f CO 2 up to 1000 µatm <strong>and</strong> CO 2 flux ~3.9–4.0 mol C m −2 yr −1 (Torres et al.2003 <strong>and</strong> references <strong>the</strong>rein). These authors concluded that CO 2 outflux is a highly variable shorttermprocess, depending on <strong>the</strong> intensity <strong>of</strong> <strong>the</strong> wind-driven upwelling, <strong>the</strong> depth <strong>of</strong> <strong>the</strong> upwelledwaters, <strong>the</strong> OM degradation (positively correlated with <strong>the</strong> apparent oxygen utilisation, AOU) <strong>and</strong><strong>the</strong> biological uptake. Fur<strong>the</strong>rmore, <strong>the</strong> high degradation rates <strong>of</strong> organic carbon by <strong>the</strong> microplanktoncommunity (dissolved organic carbon-, DOC; 1.1–21.6 µM h −1 ; G. Daneri unpublished data)<strong>and</strong> high respiration rates (81–481 mmol O 2 m −2 d −1 ; R.A. Quiñones unpublished data) indicate208
THE HUMBOLDT CURRENT SYSTEM OF NORTHERN AND CENTRAL CHILEthat nutrient recycling in <strong>the</strong> water column is an important process in <strong>the</strong> CO 2 production, resultingin high concentrations <strong>of</strong> CO 2 <strong>and</strong> outgassing when upwelled waters are warming up at <strong>the</strong> seasurface. A substantial proportion <strong>of</strong> <strong>the</strong> carbon assimilated by primary producers is reaching <strong>the</strong>bottoms via biogenic CaCO 3 flux, as has been observed <strong>of</strong>f Coquimbo (30°S) where sediment trapslocated at 2300 m water depth (~180 km <strong>of</strong>f <strong>the</strong> coast) revealed that almost 40–90% <strong>of</strong> carbonateflux is associated with some species <strong>of</strong> foraminiferans characteristic <strong>of</strong> upwelling areas (H.E.González et al. 2004a, Marchant et al. 2004). These authors suggest that biogenic CaCO 3 is <strong>the</strong>main pathway by which carbon is removed from <strong>the</strong> upper ocean, controlled by autochthonous <strong>and</strong>allochthonous foraminiferans, that is, large-size organisms with high sinking rates (1.5 days) <strong>and</strong>smaller organisms that are laterally advected. Therefore, part <strong>of</strong> this carbon is exported <strong>of</strong>fshore,sequestered from <strong>the</strong> water column <strong>and</strong> preserved in <strong>the</strong> sediments (Hebbeln et al. 2000a,b), but ithas not been clearly established what percentage <strong>of</strong> <strong>the</strong> total CO 2 assimilated by primary producersthis CaCO 3 flux represents.Macro- <strong>and</strong> micronutrient distributionThe distribution <strong>of</strong> nutrients shows a high variability in <strong>the</strong> water column associated with upwellingpulses <strong>and</strong> mixing. High concentrations occur inshore <strong>and</strong> usually decrease in <strong>the</strong> <strong>of</strong>fshore direction,followed by decreasing pigment concentrations (Escribano et al. 2003, Marín et al. 2003a, 2004a,Peñalver 2004). Off 30°S, high surface concentrations <strong>of</strong> nitrate, phosphate <strong>and</strong> silicate have beenreported (~5–15, 0.5–1 <strong>and</strong> 5–8 µM, respectively; Peñalver 2004). The nutrient distribution at thislatitude is also affected by <strong>the</strong> topography <strong>of</strong> <strong>the</strong> area, where several isl<strong>and</strong>s reduce <strong>the</strong> circulation<strong>and</strong> mixing <strong>of</strong> <strong>the</strong> water column (Peñalver 2004). In nor<strong>the</strong>rn Chile, between 20°S <strong>and</strong> 22°S, highconcentrations <strong>of</strong> nitrate were also observed near shore at 2 µM; Morales et al. 1996) <strong>and</strong> coincident with low oxygen concentrations(1000 mm yr −1 ; www.meteo<strong>chile</strong>.cl).In general, rivers play an important role in <strong>the</strong> fluxes <strong>of</strong> trace metals, nutrient <strong>and</strong> particulate matterto coastal waters, <strong>and</strong> some <strong>of</strong> <strong>the</strong>se components are considered to be important factors determiningPP in <strong>the</strong> water column. For example, Fe has been proposed as a factor limiting PP in watersenriched with o<strong>the</strong>r macronutrients but low in pigment concentrations (Martin & Gordon 1988,Martin et al. 1993, Coale et al. 1998). Some recent studies suggest that this element should berelevant in PP <strong>of</strong>f nor<strong>the</strong>rn Chile, where low dissolved Fe concentrations have been measured(0.6–1.4 nM; R. Torres unpublished data). Additionally, o<strong>the</strong>r elements such as Cd <strong>and</strong> Co mayalso play an important role in biological processes. The concentration <strong>of</strong> dissolved Co in <strong>the</strong> watercolumn shows a similar vertical distribution as macronutrients in an upwelling region <strong>of</strong>f Peru(8–10°S), apparently controlled by biological uptake <strong>and</strong> remineralisation (Saito 2005). In <strong>the</strong> samesense, dissolved Cd in <strong>the</strong> water column for Mejillones Bay shows a typical micronutrient-likedistribution (23°S; J. Valdés unpublished data). Low values <strong>of</strong> dissolved Cd (~0.4–1.6 nM) werefound in <strong>the</strong> water column, while very high concentrations were measured in surface sediments(~60 µg g –1 ; Valdés et al. 2003). In o<strong>the</strong>r coastal waters <strong>of</strong> nor<strong>the</strong>rn-<strong>central</strong> Chile, high Cd valuesin sediments are probably associated with biological uptake <strong>and</strong> subsequent deposition in <strong>the</strong>209