LETTERS PUBLISHED ONLINE: 30 MAY 2016 | DOI: 10.1038/NGEO2722 Partial decoupling of primary productivity from upwelling in the California Current system Lionel Renault1*, Curtis Deutsch2, James C. McWilliams1, Hartmut Frenzel2, Jun-Hong Liang3,4 and François Colas5 Coastal winds and upwelling of deep nutrient-rich water along subtropical eastern boundaries yield some of the ocean’s most productive ecosystems1 . Simple indices of coastal wind strength have been extensively used to estimate the timing and magnitude of biological productivity on seasonal and interannual timescales2 and underlie the prediction that anthropogenic climate warming will increase the productivity by making coastal winds stronger3–6 . The effect of wind patterns on regional net primary productivity is not captured by such indices and is poorly understood. Here we present evidence, using a realistic model of the California Current system and satellite measurements, that the observed slackening of the winds near the coast has little effect on near-shore phytoplankton productivity despite a large reduction in upwelling velocity. On the regional scale the wind drop-off leads to substantially higher production even when the total upwelling rate remains the same. This partial decoupling of productivity from upwelling results from the impact of wind patterns on alongshore currents and the eddies they generate. Our results imply that productivity in eastern boundary upwelling systems will be better predicted from indices of the coastal wind that account for its offshore structure. Upwelling indices are based on a large-scale pressure-gradient estimate of the wind field, but the spatial structure of the surface winds in eastern boundary upwelling systems (EBUS) is complex, and so is the oceanic response. Alongshore winds are typically strongest offshore, becoming weaker towards the coast owing to orography, surface roughness, and air–sea interaction7 . The nearshore drop-off in winds diminishes coastal upwelling, spreading it over a broader offshore region with slower vertical velocities (‘Ekman pumping’). It can also modulate the mean current structure8 . The partition of the total wind-driven upwelling between rapid coastal and slower offshore components has been suggested to influence the upper trophic levels of the ecosystem9,10 . However, the impact of the wind drop-off on mesoscale activity and total net primary productivity (NPP) at a regional scale has not yet been assessed11 . To investigate the influence of the coastal wind drop-off on NPP, we conducted simulations of the California Current system with an oceanic circulation and biogeochemical model (see Supplementary Methods). The model is forced by realistic climatological surface and open boundary conditions in three simulations that differ only in the cross-shore gradient of alongshore wind, the main component of mean wind stress curl (Fig. 1a). A base case (‘uniform’) is constructed from satellite scatterometer wind data, with a simple extrapolation from its reliable offshore measurements across its blind zone to the shoreline11 . Two additional simulations are conducted with wind stress reduced by 60% at the coast, consistent with observations for the upwelling season, that is, spring and summer7 . The cross-shore wind tapering distance is applied over widths of 25 km (‘sharp’) and 80 km (‘wide’) (Fig. 1a), which span the variation of the drop-off scale7 . Neither ‘sharp’ nor ‘wide’ profiles can be considered the most realistic because the real drop-off scale is not uniform7 . Model solutions are analysed along the central California coast, between 38◦ N and 43◦ N and within 100 km from shore, where the alongshore wind, eddy kinetic energy (EKE), and biological productivity are all relatively high (see Supplementary Fig. 1). Analyses are carried out for the spring season because the reversal of coastal winds during this season initiates the phytoplankton bloom timing12,13 , and an accumulation of surface nutrients during that season ensures that higher productivity persists into summer. Consistent with Ekman theory, a stronger drop-off diminishes the horizontal transport of surface water and thus the upwelling into the photic zone near the coast (Fig. 1b). Despite the weaker coastal upwelling, rates of NPP integrated over the photic zone (0–70 m, Fig. 2d) and within 20 km of the coast do not decrease and even slightly increase (Fig. 1c). In all cases, horizontal transport is constrained by the same wind stress at 100 km, so that offshore Ekman pumping compensates for differences in coastal upwelling, to maintain a similar total upwelling mass flux. Nevertheless, the integrated NPP over the photic zone (0–70 m) significantly increases by 30% when using a broader wind drop-off (‘wide’, Fig. 1c). The increases in NPP are even larger for regions closer to the shore (a 36% increase from 40 km) and in surface waters (0–10 m: a 75% increase from 0–100 km; not shown). These results imply that despite being limited by N supply, NPP is not simply related to the strength of wind-driven upwelling, either at the coast or on a broader regional scale. To identify the oceanic wind response that decouples NPP from upwelling rates, we computed the nutrient budget of the photic zone (0–70 m) within 100 km of the shore along the central California coast (38◦ –43◦ N). Although the model accounts for multiple potentially limiting nutrients, the reservoir and flux of nitrate (NO3 − ) are what limit overall production in the photic zone. Its budget can be expressed (see Supplementary Methods) as: ∂N = Fmean + Feddy − J (N ) ∂t 1 Department of Atmospheric and Oceanic Sciences, UCLA, 405 Hilgard Avenue, California 90095-1565, USA. 2 University of Washington, School of Oceanography, Box 357940 Seattle, Washington 98195-7940, USA. 3 Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA. 4 Center for Computation and Technology, Louisiana State University, Baton Rouge, Louisiana 70803, USA. 5 Institut de Recherche pour le développement (IRD), UMR LOCEAN, IRD/Sorbonne Universités (UPMC Univ Paris 06)/CNRS/MNHN, 4 Place Jussieu, Paris Cedex 75252, France. *e-mail: [email protected] NATURE GEOSCIENCE | VOL 9 | JULY 2016 | www.nature.com/naturegeoscience © 2016 Macmillan Publishers Limited. All rights reserved 505 NATURE GEOSCIENCE DOI: 10.1038/NGEO2722 LETTERS a Wind factor Factor 1.0 0.5 0.0 −100 Vertical velocity (m d−1) b 8 6 4 Uniform Sharp Wide −80 −60 −40 −20 Cross-shore distance, d (km) 0 20 0 20 0 20 Vertical velocities Uniform: 9.4 × 104 m2 d−1 Sharp: 9.8 × 104 m2 d−1 Wide: 9.2 × 104 m2 d−1 2 0 −2 −100 −80 c −60 −40 −20 Cross-shore distance, d (km) NPP (102 mmol m−1 s−1) NPP 15 10 Uniform: 5.54 × 107 mmol s−1 Sharp: 6.32 × 107 mmol s−1 Wide: 7.22 × 107 mmol s−1 5 −100 −80 −60 −40 −20 Cross-shore distance, d (km) Figure 1 | Impact of wind drop-off on total upwelling and NPP. a, Coastal wind profile factor applied to the wind product. b, Mean vertical velocity at 70 m depth between 38◦ N and 43◦ N during spring (April–June). The total upwelling rate integrated over a distance of 100 km offshore is indicated in the legend. c, NPP integrated over the photic zone (0–70 m depth) and between 38◦ N and 43◦ N during spring. The total NPP (integrated over a distance of 100 km offshore) is indicated in the legend. The shaded areas represent the standard deviations. The means and standard deviations are estimated using 8 years of simulations. where N represents nitrate concentration and (∂N /∂t) is the buildup of nitrate inventory change over time according to the imbalance between total physical nitrate transport, due to both the time-averaged flow (Fmean ) and its fluctuations (Feddy ), and the total nitrate uptake by the ecosystem, J (N ). The biological uptake of nitrate approximates the net community production of organic nitrogen (dissolved and particulate) that is exported to depth. During spring, when winds become upwelling favourable, the physical nitrate supply (Fmean + Feddy ) exceeds the rate of ecosystem production in all cases, leading to a significant buildup of the nitrate reservoir (Fig. 2a). The larger integrated NPP in cases with a wind drop-off (‘sharp’ and ‘wide’) is reflected in J (N ) and the more rapid accumulation of surface nutrients (∂N /∂t) ensures that higher productivity persists into summer. In the case of a broader wind drop-off (‘sharp’ and ‘wide’), both the increase of J (N ) and (∂N /∂t) is due to the higher physical nutrient delivery. As the boundary layer is shallower than 70 m (estimated from the K-profile parameterization, not shown), the effect of wind shape on nutrient supply must be due to changes in advective transport rather than mixing. A broader wind drop-off alters significantly the mean alongshore currents, especially the coastal undercurrent, which transports 506 high-nutrient tropical water poleward. Consistent with Sverdrup dynamics, a stronger wind stress curl in ‘sharp’ and ‘wide’ yields a stronger, shallower poleward flow8 that induces a weaker mean southward flow between 0–70 m depth (Fig. 2b), and a higher nitrate transport than in ‘uniform’. For the coastal region (0–20 km), the reduction of the vertical velocity from ‘uniform’ to ‘wide’ (by 54%, Fig. 1b) is offset by the stronger alongshore undercurrent that brings higher nutrient water below the photic layer. However, the increase of nitrate transport does not continue from ‘sharp’ to ‘wide’, in spite of a further strengthening and shoaling of the mean undercurrent. Thus, the change in the mean undercurrent only partially explains the larger advective supply rate induced by a broader wind drop-off. Moreover, the effect of wind shape on the alongshore nutrient supply is offset by changes in nutrient transport to the east, such that horizontal nutrient fluxes in the photic layer as a whole are insensitive to the wind drop-off (Fig. 2a). Hence, the nitrate supply by mean vertical velocities is also roughly insensitive to the wind drop-off (Supplementary Fig. 8). Together, these results imply a major role for nutrient transport by fluctuating components of the circulation. Although the wind structure has a relatively small effect on Fmean , it exerts a strong indirect influence on the mesoscale eddies that also have a net nutrient transport (Feddy ). A broader wind drop-off in ‘sharp’ and ‘wide’ weakens the vertical shear of the alongshore current below the thermocline, flattens the isopycnal tilt, and reduces the EKE (Fig. 2c). The rate at which EKE is converted from eddy potential energy by baroclinic instability is diagnosed from the integrated eddy vertical buoyancy flux, which is reduced under the wind drop-off (Supplementary Fig. 6). In the EBUS, eddies have been shown to reduce NPP by subducting nutrients along isopycnal surfaces that plunge below the euphotic layer offshore, termed ‘eddy quenching’14 . Indeed, all our cases show a negative (downward) eddy nitrate flux during spring (Fig. 2d), except in the first 10 km nearshore, where the upwelling prevails. By diminishing the mesoscale eddy activity, a broader wind drop-off in ‘sharp’ and ‘wide’ weakens significantly this removal of nitrate from the photic zone and thus allows its more complete consumption. This inverse relationship between ‘eddy quenching’ and the wind drop-off accounts for the minimal response of NPP to winds near the coast, and the shallow subduction of nitrate contributes to the higher NPP offshore under a strong wind drop-off14,15 . Ekman theory and model simulations predict that the wind stress magnitude is the main driver of productivity and, further, that wind drop-off modulates NPP by being negatively correlated with EKE, but positively correlated with phytoplankton growth and biomass. We tested these predictions using satellite data for chlorophyll a (a proxy for NPP, from SeaWiFS), wind stress (from QuikSCAT), wind stress curl (a measure of the wind drop-off), and the EKE (from the Archiving, Validation, and Interpretation of Satellite Oceanographic data (AVISO)) (Fig. 3a–c). The scatterometer blind zone near the coast allows only a partial sampling of the wind drop-off profile16 ; therefore, a positive wind curl anomaly can be interpreted as a broader wind drop-off more fully sampled by QuikSCAT, and a negative anomaly implies a sharper wind drop-off, more of which occurs within the blind zone. The offshore wind stress gives a reliable metric for the total upwelling. Over the period of overlapping satellite records (2000–2009), interannual fluctuations in the wind stress, wind stress curl, EKE and chlorophyll are significantly correlated. Consistent with earlier studies17 , the mean upwelling is the main driver of the productivity. However, years with a larger wind stress curl generally have smaller EKE and larger chlorophyll a (Fig. 3d). The former link implicates eddy modulation by wind-induced changes in the unstable alongshore currents, and the latter link supports eddy quenching of NPP. The importance of the wind drop-off in modulating total NPP is most evident in years when anomalies NATURE GEOSCIENCE | VOL 9 | JULY 2016 | www.nature.com/naturegeoscience © 2016 Macmillan Publishers Limited. All rights reserved NATURE GEOSCIENCE DOI: 10.1038/NGEO2722 a 20 Rate (106 mmol s−1) 15 LETTERS Nutrient budget Uniform Sharp Wide Horizontal transport Vertical transport 10 5 0 −5 −10 Storage b Uptake Transport Mean c Uniform Sharp Wide d Uniform Sharp Wide 44 40 38 42 Depth (m) Latitude (° N) 42 40 38 36 36 −0.2 0 0.2 Alongshore current (m s−1) Total 0 100 200 300 400 EKE (cm2 s−2) Mean Eddy Sharp Uniform −20 −40 −60 −80 −100 −120 −140 −160 −180 −200 −100 Total Wide 15 10 5 0 −5 (mmol m−2 s−1) Latitude (° N) 44 Eddy −10 −50 d (km) 0 −100 −50 d (km) 0 −100 −50 d (km) 0 −15 Figure 2 | Wind drop-off control of the NPP by modulation of the eddy physical fluxes. a, Nutrient budget during spring between 38◦ N and 43◦ N from 70 m depth to the surface from the three model experiments. Storage is ∂N/∂t. Uptake is J(N). Transport is F. b, Alongshore current during spring, averaged over a cross-shore band 100 km wide and 70 m in depth. The dashed black lines indicate the 38◦ –43◦ N region. c, Mean surface EKE, averaged over a cross-shore band 100 km wide. The dashed black lines indicate the 38◦ –43◦ N region. d, Mean vertical eddy nutrient supply during spring averaged between 38◦ N to 43◦ N. The black lines indicate the corresponding mean simulated euphotic depth. d, cross-shore distance. The error bars (a) and shaded areas (b,c) represent the standard deviations estimated using 8 years of simulation. 33 30 129 126 123 120 117 Longitude (° W) c 150 42 100 39 36 50 Latitude (° N) 36 45 3.00 42 1.00 0.60 39 0.20 0.10 36 0.05 33 33 30 45 129 126 123 120 117 Longitude (° W) d 30 0 Chl-a (mg m−3) 39 b EKE (cm2 s−2) Latitude (° N) 42 10 8 6 4 2 0 −2 −4 −6 −8 −10 Latitude (° N) 45 Wind stress curl (N m−2 per 104 km) a 129 126 123 120 117 Longitude (° W) Indices Class 1: reinforcing processes Class 2: counteracting processes 1 0 −1 2001 2002 2006 2004 2005 2007 Year 2008 2009 2000 2003 Wind index Curl index EKE index Chl-a index c(Wind, Curl) = 0.1 c(Wind, EKE) = −0.2 c(Wind, Chl-a) = 0.6 c(Curl, EKE) = −0.5 c(Curl, Chl-a) = 0.3 c(EKE, Chl-a) = −0.3 Figure 3 | An upwelling index that considers wind structure, and perhaps eddy activity, would better predict interannual NPP variations. a, Mean wind stress curl from QuikSCAT during spring. Mean wind stress magnitude is superimposed with dashed black contours. The solid black contour indicates the location where the indices and the budget shown in Fig. 2 are computed. b, EKE from AVISO. c, Chlorophyll a (Chl-a) from SeaWiFS. d, Indices of variability in wind stress, wind stress curl, EKE and chlorophyll a. The indices are computed by subtracting the mean value over the area indicated by the solid black contour in a–c from 2000–2009 in spring, and dividing the resulting anomalies by the largest magnitude over the time period. The correlations among the indices are listed, and they all are significant at the 95% level. The years are divided into two categories: when the mean upwelling and wind drop-off effect are reinforcing; and when they are counteracting. Similar results are found for the summer season. NATURE GEOSCIENCE | VOL 9 | JULY 2016 | www.nature.com/naturegeoscience © 2016 Macmillan Publishers Limited. All rights reserved 507 NATURE GEOSCIENCE DOI: 10.1038/NGEO2722 LETTERS in wind stress curl counteract, rather than reinforce, the changes in coastal upwelling (Fig. 3d). Such counteracting years occur in roughly half the years in the available time series, when stronger coastal winds are associated with weaker wind stress curl, or vice versa. In three out of four such years in the satellite record, the NPP anomaly is in the direction predicted by the wind drop-off, and opposite to what would be expected by the anomaly in coastal wind. Satellite estimates thus are consistent with the model predictions and suggest that relatively complex indices, not solely based on simple wind time series, will be needed to predict interannual NPP variations in eastern boundary upwelling systems. This consistency between satellite remote sensing and regional modelling experiments supports a new eddy-mediated link between the coastal wind pattern and biological productivity, at least for the California Current system. The same mechanisms are likely to be present in other EBUS, albeit to varying degrees that reflect differences in wind structure, ocean stratification, and nutrient limitation factors. For example, the steeper coastal orography of the Andes in South America may induce a broader wind drop-off7 , which may explain the weak nearshore generation of EKE18 , thus yielding less eddy quenching of nutrients and a more productive system17 . Similarly, the density stratification has a strong influence on baroclinic energy conversion and EKE levels, so the wind dropoff effect can only partially explain the EKE difference between the EBUS. As the mean upwelling is the main driver of the productivity, indices based on large-scale winds remain useful to predict the overall tendencies of coastal marine productivity. However, our findings help explain residual interannual variations of NPP in the EBUS and demonstrate the need for better predictors than indices based on large-scale winds alone19 . Predicting how productivity in EBUS will react to future climate change will require regional atmospheric and/or coupled models that adequately resolve the wind drop-off profile and the ocean–atmosphere interactions20 , as well as changes in the oceanic state that modulate its effects on eddydriven nutrient supply. Methods Methods, including statements of data availability and any associated accession codes and references, are available in the online version of this paper. Received 6 August 2015; accepted 25 April 2016; published online 30 May 2016 References 1. Carr, M. E. & Kearns, E. J. Production regimes in four Eastern Boundary Current systems. Deep-Sea Res. II 50, 3199–3221 (2003). 2. Bograd, S. J. et al. Phenology of coastal upwelling in the California Current. Geophys. Res. Lett. 36, 1602 (2009). 3. Bakun, A. Global climate change and intensification of coastal ocean upwelling. Science 247, 198–201 (1991). 4. Wang, D., Gouhier, T. C., Menge, B. A. & Ganguly, A. R. Intensification and spatial homogenization of coastal upwelling under climate change. Nature 518, 390–394 (2015). 508 5. Sydeman, W. J. et al. Climate change and wind intensification in coastal upwelling ecosystems. Science 345, 77–80 (2014). 6. Bakun, A. et al. Anticipated effects of climate change on coastal upwelling ecosystems. Curr. Clim. Change Rep. 1, 85–93 (2015). 7. Renault, L., Hall, A. & McWilliams, J. C. Orographic shaping of US West Coast wind profiles during the upwelling season. Clim. Dynam. 46, 273–289 (2016). 8. Song, H., Miller, A. J., Cornuelle, B. D. & Di Lorenzo, E. Changes in upwelling and its water sources in the California Current System driven by different wind forcing. Dyn. Atmos. Oceans 52, 170–191 (2011). 9. Rykaczewski, R. R. & Checkley, D. M. Influence of ocean winds on the pelagic ecosystem in upwelling regions. Proc. Natl Acad. Sci. USA 105, 1965–1970 (2008). 10. Jacox, M. G., Moore, A. M., Edwards, C. A. & Fiechter, J. Spatially resolved upwelling in the California Current System and its connections to climate variability. Geophys. Res. Lett. 41, 3189–3196 (2008). 11. Capet, X. J., Marchesiello, P. & McWilliams, J. C. Upwelling response to coastal wind profiles. Geophys. Res. Lett. 31, 13 (2004). 12. Barth, J. A. et al. Delayed upwelling alters nearshore coastal ocean ecosystems in the northern California current. Proc. Natl Acad. Sci. USA 104, 3719–3724 (2007). 13. Thomas, A. C. & Brickley, P. Satellite measurements of chlorophyll distribution during spring 2005 in the California Current. Geophys. Res. Lett. 33, L22S05 (2006). 14. Gruber, N. et al. Eddy-induced reduction of biological production in eastern boundary upwelling systems. Nature Geosci. 4, 787–792 (2011). 15. Nagai, T. et al. Dominant role of eddies and filaments in the offshore transport of carbon and nutrients in the California Current System. J. Geophys. Res. 120, 5318–5341 (2015). 16. Renault, L. et al. Impact of atmospheric coastal jet off central Chile on sea surface temperature from satellite observations (2000–2007). J. Geophys. Res. 114, C08006 (2009). 17. Chavez, F. P. & Messié, M. A comparison of eastern boundary upwelling ecosystems. Prog. Oceanogr. 83, 80–96 (2009). 18. Colas, F., McWilliams, J. C., Capet, X. & Kurian, J. Heat balance and eddies in the Peru-Chile current system. Clim. Dynam. 39, 509–529 (2012). 19. García-Reyes, M., Largier, J. L. & Sydeman, W. J. Synoptic-scale upwelling indices and predictions of phyto- and zooplankton populations. Prog. Oceanogr. 120, 177–188 (2014). 20. Renault, L. et al. Modulation of wind-work by oceanic current interaction with the atmosphere. J. Phys. Oceanogr. 46, 1685–1704 (2016). Acknowledgements We appreciate support from the Office of Naval Research (N00014-12-1-0939), National Science Foundation (OCE-1419450 and OCE-1419323), Bureau of Ocean Energy Management, and California Ocean Protection Council, as well as computing resources from the Extreme Science and Engineering Discovery Environment and on the Yellowstone cluster (ark:/85065/d7wd3xhc) provided by NCAR’s Computational and Information Systems Laboratory, sponsored by the National Science Foundation. Author contributions L.R., J.C.M. and C.D. conceived and designed the experiments; L.R. performed the experiments; L.R., C.D., J.C.M., H.F. and F.C., analysed the data; L.R., H.F. and J.-H.L. contributed materials/analysis tools; L.R., C.D. and J.C.M. co-wrote the paper. Additional information Supplementary information is available in the online version of the paper. Reprints and permissions information is available online at www.nature.com/reprints. Correspondence and requests for materials should be addressed to L.R. Competing financial interests The authors declare no competing financial interests. NATURE GEOSCIENCE | VOL 9 | JULY 2016 | www.nature.com/naturegeoscience © 2016 Macmillan Publishers Limited. All rights reserved NATURE GEOSCIENCE DOI: 10.1038/NGEO2722 LETTERS Methods Model configuration. The oceanic simulations were performed with the Regional Oceanic Modeling System (ROMS)21 . ROMS is a free-surface, terrain-following coordinate model with split-explicit time stepping and Boussinesq and hydrostatic approximations. The model extends from 142.1◦ W to 114.4◦ W and from 23.9◦ N to 50.0◦ N (see Supplementary Fig. 1). The model grid has 627 × 377 points with a horizontal resolution of 4 km and has 42 vertical levels. The vertical grid is stretched for increased resolution of the surface and bottom boundary layers. The bottom topography is derived from an SRTM30 database22 . The boundary condition algorithm consists of a modified Flather-type scheme for the barotropic mode23 and Orlanski-type scheme for the baroclinic mode (including temperature and salinity; ref. 24). The simulation is forced at the surface by the QuikSCAT-based daily product described in ref. 25 (based on the SCOW climatology). Heat and freshwater atmospheric forcing are from the Comprehensive Ocean–Atmosphere Data Set26 . The freshwater atmospheric forcing has an additional restoring term to prevent surface salinity from drifting away from climatological values. This weak restoring is towards climatological monthly surface salinity from the World Ocean Atlas27 . A flux correction term is included in the atmospheric heat forcing to allow feedback from the ocean to the atmosphere following the formulation of ref. 28. As in ref. 29, initial and boundary information are taken from a 12 km Pacific climatological solution and the model is run for ten years. Ref. 25 has full information about a similar Pacific simulation at coarser resolution. The Biogeochemical Elemental Cycling30 model is coupled to ROMS. It includes multiple limiting nutrients (N, P, Fe and Si) and three phytoplankton functional groups (diatoms, diazotrophs, and small phytoplankton) that represent the biogeographical variability of different oceanic biomes, for example, highly productive coastal regimes versus the oligotrophic open ocean areas of the subtropical gyres. It includes the dissolved iron cycle, including inputs of iron from sediments and from atmospheric dust deposition. The degree of realism of the simulation here is similar to the results of ref. 14. A set of three experiments has been carried out. The only difference between them is the coastal wind profile used to force the model (Fig. 1a). ‘uniform’ is the control run, the QuikSCAT SCOW wind is interpolated onto the ROMS grid, and the missing coastal values are tapered using a simple extrapolation11 . As QuikSCAT monitors only partially the wind drop-off in this region, we consider this experiment as without wind drop-off. ‘sharp’ and ‘wide’ add a wind drop-off using the factors shown in Fig. 1a. As a result, ‘sharp’ and ‘wide’ have wind reduction by 60% and a wind drop-off length of 25 km and 80 km, respectively. Note, as shown by ref. 7, the wind drop-off is not uniform and presents latitudinal variation both in length and wind reduction. The values chosen for this study are in the range of values found in that former study. However, for the purpose of this study, idealized experiments using such a wind modification allow us to assess how the coastal wind shape controls mesoscale activity and NPP. In this study, the winter, spring, summer and autumn seasons correspond to the months (January–March), (April– June), (July–September) and (October–December), respectively. Budget analysis. To analyse the role of eddies in the nutrient evolution, we decompose the advective flux of any nutrient concentration C into time-mean and fluctuation (eddy) components: UC = Ū C̄ + U 0 C 0 (1) where U denotes the three-dimensional velocity. As in ref. 31, where an analysis is made for C equal to the buoyancy, the ‘instantaneous’ simulation outputs are 2-day averages, and the mean is defined as a multi-year seasonal average. In the following we denote the alongshore and cross-shore currents as v and u and the offshore distance as d. The balance equation for any C is ∂C = ∇ · K · ∇(C) − ∇h · uh C − ∇v · wC − J (C) ∂t (2) where K is the eddy kinematic diffusivity tensor, ∇ is the three-dimensional gradient operators, and uh , and w are the horizontal and vertical velocities, and J (C) is the biogeochemical source minus sink term. We present the budget for inorganic nitrogen, because it is the nutrient that ultimately limits biological productivity at a regional scale in the California Current. We further restrict our analysis to nitrate (NO3 − ), because it is by far the largest reservoir and physical supply of inorganic nitrogen (compared with ammonium transport and concentration, which are both very small). Nitrate also has the advantage of not being sensitive to the internal ecosystem nutrient transformations. The only sinks of nitrate are from phytoplankton uptake, and its only biological source is nitrification, which is inhibited by light in the photic zone. The biological uptake of nitrate in the model is thus equal to ‘new production’, and balances the portion of NPP that is exported to depth. From equation (2) a mean-seasonal balance is estimated for the 38◦ –43◦ N × 100 km cross-shore × 70 m depth region, which includes both the turbulent surface boundary layer and euphotic zone in spring. By 70 m depth the mixing term becomes negligible. Equation (2) will be first analysed in the reduced form, dN = Ftot − J (N ) dt (3) where N represents nitrate concentration, Ftot is the total physical nitrate transport, and J (N ) the total nitrate uptake by the ecosystem. This budget analysis is then further decomposed between mean transport (Fmean ) and eddies (Feddies ) following equation (1), ∂N = Fmean + Feddy − J (N ) (4) ∂t and between horizontal (Fhor ) and vertical transport (Fver ), ∂N = Fhor(mean) + Fhor(eddy) + Fver(mean) + Fver(eddy) − J (N ) ∂t (5) Data availability and description. QuikSCAT wind stress. The near-surface atmospheric circulation over the ocean is described through daily QuikSCAT zonal and meridional wind components, obtained from Centre ERS d’Archivage et de Traitement on a 0.25◦ × 0.25◦ resolution grid32 . This product is built from both ascending and descending passes from discrete observations (available in JPL/PO.DAAC Level 2B product) over each day. Standard errors are also computed and provided as complementary gridded fields. SeaWiFS chlorophyll a. Surface chlorophyll concentrations were estimated from SeaWiFS data33 for the 2000–2009 period. We used Level 3 (9 km) monthly composites obtained from the Distributed Active Archive Center at NASA Goddard Space Flight Center. AVISO sea level anomalies. The sea level anomalies come from the Archiving, Validation, and Interpretation of Satellite Oceanographic data (AVISO) multimission mapped altimetry product34 . We use the Delayed Time 2014 version, in which data from at least two (up to four) simultaneous satellite altimeter missions were merged and mapped onto a 0.25◦ Mercator grid at daily intervals for the period October 1992–December 2013; the 1993–1999 mean was removed at each grid point. The surface geostrophic currents are computed by using the sea level anomalies and the eddy kinetic energy (EKE) is defined as 1/2(u02 + v 02 ) where u0 and v 0 are velocity perturbations relative to a seasonal time-mean (same method is applied when estimating the EKE from the model). California Cooperative Oceanic Fisheries Investigation. Large-scale systematic hydrographic sampling of the California Current system was initiated in 1949 by the California Cooperative Oceanic Fisheries Investigations (CalCOFI) programme. Since 1950, stations have been repeatedly occupied at varying intervals based on a geographically fixed grid. In this study, lines 60 (off Point Reyes; ∼38◦ N) and 67 (∼37◦ N), which have enough data to estimate a seasonal climatology of respectively chlorophyll a and temperature, are used to validate the simulations. Satellite analysis. Indices of wind stress, wind stress curl, EKE and chlorophyll a as estimated over the black rectangle in Fig. 3 during spring are computed using the following method: the long-time mean value over the black rectangle during spring over the period 2000–2009 is calculated first. Then, the indices are calculated by computing the anomalies of spring rectangle values with respect to the long-time mean value and are finally normalized by the largest magnitude over the time period. Mean variability of chlorophyll a, wind stress and EKE. Model solutions are analysed along the central California coast during spring, between 38◦ N and 43◦ N and within 100 km from shore. The spring season is chosen because the reversal of coastal winds during this season initiates the phytoplankton bloom timing and an accumulation of surface nutrients during that season will ensure that higher productivity persists into summer. Additionally, during spring, as illustrated in Supplementary Fig. 1, the alongshore wind stress, EKE and biological productivity are all relatively high. Model evaluation and eddy buoyancy fluxes. To illustrate the realism of the simulations, Supplementary Figs 2–6 depict some basic diagnostics of both physical and biogeochemical fields. Supplementary Fig. 2 shows the sea surface temperature mean from ‘uniform’ and in situ observations (World Ocean Atlas27 ). The simulated sea surface temperature mean and variability are fairly reproduced by the model, which clearly shows the upwelling signature. Supplementary Fig. 3 represents simulated and observed vertical distributions of temperature during spring along CalCOFI line 67 (which starts around 37◦ N). It indicates that the simulated vertical gradients are captured well. However, the onshore slope is slightly overestimated, particularly in the nearshore region (Supplementary Fig. 2). Supplementary Fig. 4a shows the EKE computed from ‘uniform’ using low-pass NATURE GEOSCIENCE | www.nature.com/naturegeoscience © 2016 Macmillan Publishers Limited. All rights reserved NATURE GEOSCIENCE DOI: 10.1038/NGEO2722 LETTERS filtered (7-day averaging and Gaussian spatial filter with 30-km half-width) geostrophic velocities. The realism of the EKE indicates the ability of the model to reproduce the mesoscale activity (AVISO EKE is shown in Supplementary Fig. 4b) but also the mean current because mesoscale eddies arise from mean currents instabilities. Recently, ref. 20 showed that current feedback to the atmosphere dampens the EKE. Here, the offshore EKE is overestimated. The absence of current feedback in the model (fluxed-forced) induces an overestimation of the eddy life, allowing eddies to propagate further offshore. Supplementary Fig. 5 depicts the mean chlorophyll a from ‘uniform’ and SeaWiFS during spring. As expected, the coastal upwelling region is marked by high concentrations of chlorophyll a. Model-simulated chlorophyll a and the observations have similar agreements and disagreements as in ref. 14. There is an overall tendency for the model to be biased low. The largest underestimation occurs in the nearshore areas. Offshore, a low bias is found that is likely to be due to the absence of picoplankton (which grow under oligotrophic conditions) in the model. Finally, Supplementary Fig. 6 shows simulated (from ‘uniform’ and ‘sharp’) and observed vertical distributions of chlorophyll a during spring along CalCOFI line 60 (starting around 38◦ N). ‘uniform’ underestimates the coastal chlorophyll a concentration. By using a broader wind drop-off (here ‘sharp’, a similar increase is found in ‘wide’), the chlorophyll a concentration increases (see main paper), becoming more realistic with respect to the observations. This also illustrates the sensitivity of the simulated chlorophyll a to the coastal wind shapes. A broader wind drop-off diminishes the southward surface current, strengthens the undercurrent (Fig. 2b), and even can induce a surfacing of the undercurrent in ‘wide’ (not shown). This is consistent with Sverdrup dynamics in response to wind drop-off: a positive wind stress curl produces a barotropic poleward flow that adds up to the coastal undercurrent35,36 . As a result, the undercurrent strength is larger with a broader wind drop-off. Note, the surfacing of the undercurrent in ‘wide’ is not realistic (not shown); an overestimation of the wind drop-off length can induce such a feature. A broader wind drop-off not only changes the undercurrent depth and intensity but also induces a different vertical shear of the alongshore current. From ‘uniform’ to ‘wide’, the vertical shear diminishes below the thermocline, stabilizing the water column. This is confirmed by Supplementary Fig. 7, which depicts the mean vertical buoyancy flux w 0 b0 from all of the experiments during spring. Negative values are important as they indicate regions where eddies act locally contrary to the baroclinic instability theoretical expectation of positive conversion of available potential to kinetic energy. In the stratified interior, eddy buoyancy flux acts to balance the effect of upwelling-favourable winds, that is, to flatten the tilted upper thermocline31 . As a result, the offshore eddy restratification flux weakens in the progression from ‘uniform’ to ‘wide’, and there is a similar weakening in the eddy destratification flux near the coast. By inducing a weaker vertical shear of the alongshore current, a broader drop-off weakens the energy flux associated with upper ocean baroclinic instability and then reduces the EKE (Fig. 2c). A broader wind drop-off, by reducing the coastal wind stress, also weakens the coastal wind work35 , acting again towards a reduction of the EKE. Supplementary Fig. 8 depicts a dissolved inorganic nitrogen budget of the photic zone (0–70 m) within 20 km of the shore along the central California coast ◦ (38 –43◦ N) during spring (N , nitrate concentration). The effect of the reduction of the vertical velocities from ‘uniform’ to ‘wide’ (by 54%, Fig. 1b) is damped by the effect of changes of alongshore current on the nitrate reservoir below the photic layer. ‘wide’ leads to a more effective coastal mean upwelling because the mean nitrate supply by mean vertical and horizontal velocities is roughly insensitive to the wind drop-off (Supplementary Fig. 8), whereas the opposing eddy flux is decreased. Code availability. We have opted not to make the computer code associated with this paper available because we use in-house versions of ROMS at UCLA and UW; similar ROMS versions are available through Rutgers (www.myroms.org) and ROMS-AGRIF (http://www.romsagrif.org). Data availability. The simulation outputs that support the findings of this study are available on request from the corresponding author (L.R.). The data are not publicly available owing to the large size of the model output files. References 21. Shchepetkin, A. F. & McWilliams, J. 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World Ocean Atlas 2001: Objective Analyses, Data Statistics, and Figures: CD-ROM Documentation (US Department of Commerce, National Oceanic and Atmospheric Administration, National Oceanographic Data Center, Ocean Climate Laboratory, 2002). 28. Barnier, B., Siefridt, L. & Marchesiello, P. Thermal forcing for a global ocean circulation model using a three-year climatology of ECMWF analyses. J. Mar. Syst. 6, 363–380 (1995). 29. Molemaker, M. J., McWilliams, J. C. & Dewar, W. K. Submesoscale instability and generation of mesoscale anticyclones near a separation of the California Undercurrent. J. Phys. Oceanogr. 45, 613–629 (2015). 30. Moore, J. K., Doney, S. C. & Lindsay, K. Upper ocean ecosystem dynamics and iron cycling in a global three-dimensional model. Glob. Biogeochem. Cycles 18, GB4028 (2004). 31. Colas, F., Capet, X., McWilliams, J. C. & Li, Z. Mesoscale eddy buoyancy flux and eddy-induced circulation in Eastern Boundary Currents. J. Phys. Oceanogr. 43, 1073–1095 (2013). 32. 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