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Recent studies have described active nitrogen fixation in high-latitude waters, but the ecological controls on the occurrence or activity of nitrogen-fixing organisms (diazotrophs) in such systems remain unknown. Turnover rates and top-down controls are also general knowledge gaps for marine diazotrophs. We detected abundant UCYN-A (endosymbiotic nitrogen-fixing cyanobacteria) in the Gulf of Anadyr, western Bering Sea, which correlated with high dissolved iron to dissolved inorganic nitrogen ratios (Fe : DIN) due to riverine input. Growth and grazing mortality of UCYN-A sublineages were almost balanced with higher biomass-turnover rates compared to the whole phytoplankton community, indicating selective grazing of UCYN-A in nitrogen-depleted waters. Grazing rates on UCYN-A1 (small cells) were higher than for UCYN-A2/3/4 (large cells), consistent with the general size dependence of phytoplankton growth and grazing mortality. We found that Fe : DIN is a major determinant of UCYN-A abundances in high-latitude waters, where UCYN-A could make substantial contributions to plankton food-web cycling.

Compound-specific isotopic analysis of amino acids (CSIA-AA) is increasingly used in ecological and biogeochemical studies tracking the origin and fate of nitrogen (N). Its advantages include the potential for resolving finer-scale trophic dynamics than possible with standard bulk SIA and for reconstructing historical changes in the food webs of consumers from analyses of specimens in preserved sample archives. For the latter, assessing the effects of chemical preservatives on δ15NAA has been inconclusive because the conventional CSIA approach for derivatized AAs by gas chromatography-combustion-isotope ratio mass spectrometry (GC-C-IRMS) has analytical errors (0.4–1.0‰) in the range expected from chemical preservation. Here, we show improved analytical precision (0.15 ± 0.08‰) for 11 underivatized AA standards analyzed by high-pressure liquid chromatography followed by offline elemental analysis-IRMS (HPLC/EA-IRMS). Using this method, we report the first high-precision tests of preservation effects on δ15NAA in Northern Anchovy (Engraulis mordax) kept 1.5-yr in ethanol and up to 27-yr in formaldehyde. We found minimal methodological induced fractionation for eight AAs, and preservation effects on δ15N were similar regardless of duration and preservative used. Although some of the AAs differed significantly from frozen control samples (AA average +1.0 ± 0.8‰), changes in δ15N in the source AA phenylalanine and trophic position estimates were statistically insignificant. Our results are encouraging for resolving fine-scale natural variability using HPLC/EA-IRMS on chemically preserved specimens and for ultimately reconstructing biogeochemical records and trophic dynamics over long time scales.

Variability in community composition is often attributed to underlying differences in physical environments. However, predator–prey interactions can play an equally important role in structuring communities. Although environmental differences select for different species assemblages, less is known about their impacts on trait compositions. We develop a trait-based analysis of plankton communities of the southern California Current System across multiple trophic levels, from bacteria to mesozooplankton, and over a gradient of environmental conditions, from the oligotrophic open ocean to coastal upwelling. Across a factor of four differences in total community biomass, we observe similarities in the size structure along the environmental gradient, with the most pronounced departures from proportional variations in the biomasses found in the largest protists (> 40 µm). Differences in the trait distributions emerge within a small range of size groups: greater biomass contribution of larger autotrophs (> 10 µm) is observed only for the upwelling region.

We investigated variability in the C:234Th ratio of sinking particles and its relationship to changing water column characteristics and plankton ecological dynamics during 29 Lagrangian experiments conducted on six cruises of the California Current Ecosystem Long-Term Ecological Research (CCE-LTER) Program. C:234Th ratios of sinking particles collected by a surface-tethered sediment trap (C:234ThST) varied from 2.3 to 20.5 μmol C dpm−1 over a depth range of 47–150 m. C:234ThST was significantly greater (by a factor of 1.8) than C:234Th ratios of suspended >51-μm particles collected in the same water parcels with in situ pumps. C:234Th ratios of large (>200-μm) sinking particles also exceeded those of smaller sinking particles. C:234ThST decreased with depth from the base of the euphotic zone through the upper twilight zone. C:234ThST was positively correlated with several indices of ecosystem productivity including particulate organic carbon (POC) and chlorophyll (Chl) concentrations, mesozooplankton biomass, and the fraction of Chl >20-μm. Principal component analysis and multiple linear regression suggested that decaying phytoplankton blooms exhibited higher C:234ThST than actively growing blooms at similar biomass levels. C:234ThST was positively correlated with indices of the fractional contribution of fecal pellets in sediment traps when the proportion of fecal pellets was low in the traps, likely because of a correlation between mesozooplankton biomass and other indices of ecosystem productivity. However, when fecal pellets were a more important component of sinking material, C:234ThST decreased with increasing fecal pellet content. C:234ThST was also positively correlated with the Si:C ratio of sinking particles. Across the dataset (and across depths) a strong correlation was found between C:234ThST and the ratio of vertically-integrated POC to vertically-integrated total water column 234Th (vC:234Thtot). A mechanistic one-layer, two-box model of thorium sorption and desorption was invoked to explain this correlation. Two empirical models (one using vC:234Thtot; one using depth and vertically-integrated Chl) were developed to predict C:234Th ratios in this coastal upwelling biome. The former regression (log10(C:234ThST) = 0.43 × log10(vC:234Thtot) + 0.53) was found to also be a reasonable predictor for C:234ThST from diverse regions including the Southern Ocean, Sargasso Sea, Subarctic North Pacific, and Eastern Tropical North Pacific.

We used extensive ecological and biogeochemical measurements obtained from quasi-Lagrangian experiments during two California Current Ecosystem Long-Term Ecosystem Research cruises to analyze carbon fluxes between the epipelagic and mesopelagic zones using a linear inverse ecosystem model (LIEM). Measurement constraints on the model include 14C primary productivity, dilution-based microzooplankton grazing rates, gut pigment-based mesozooplankton grazing rates (on multiple zooplankton size classes), 234Th:238U disequilibrium and sediment trap measured carbon export, and metabolic requirements of micronekton, zooplankton, and bacteria. A likelihood approach (Markov Chain Monte Carlo) was used to estimate the resulting flow uncertainties from a sample of potential flux networks. Results highlight the importance of mesozooplankton active transport (i.e., diel vertical migration) in supplying the carbon demand of mesopelagic organisms and sequestering carbon dioxide from the atmosphere. In nine water parcels ranging from a coastal bloom to offshore oligotrophic conditions, mesozooplankton active transport accounted for 18–84% (median: 42%) of the total carbon transfer to the mesopelagic, with gravitational settling of POC (12–55%; median: 37%), and subduction (2–32%; median: 14%) providing the majority of the remainder. Vertically migrating zooplankton contributed to downward carbon flux through respiration and excretion at depth and via mortality losses to predatory zooplankton and mesopelagic fish (e.g., myctophids and gonostomatids). Sensitivity analyses showed that the results of the LIEM were robust to changes in nekton metabolic demand, rates of bacterial production, and mesozooplankton gross growth efficiency. This analysis suggests that prior estimates of zooplankton active transport based on conservative estimates of standard (rather than active) metabolism are likely too low.

The 2014–2015 warm anomaly (aka "the Blob"), the largest of periodic and intensifying marine heat wave (MHW) perturbations in the northeast Pacific, may provide some insight about the future warmer ocean. Here, we use mixed‐layer carbon estimates for total phytoplankton, major size classes and functional groups from 45 CalCOFI cruises to: (1) compare 2014–2015 MHW impacts in the southern California Current System to baseline estimates from 2004 to 2013 and (2) to test a space‐for‐time exchange hypothesis that links biomass structure to variability of nitracline depth (NCD). Seasonal and inshore‐offshore analyses from nine stations revealed almost uniform 2°C MHW warming extending 700 km seaward, fourfold to sixfold declines in nitrate concentration and 18‐m deeper NCDs. Phytoplankton C decreased 16–21% compared to 45–65% for Chl a, with the threefold difference due to altered C : Chl a. Among size classes, percent composition of nanoplankton decreased and picophytoplankton increased, driven by higher Prochlorococcus biomass, while Synechococcus and picoeukaryotes generally declined. Diatom and dinoflagellate C decreased in both onshore and offshore waters. Seasonally, the MHW delayed the normal winter refresh of surface nitrate, resulting in depressed stocks of total phytoplankton and nanoplankton, Synechococcus and picoeukaryotes during winter. Consistent with the space‐for‐time hypothesis, biomass variations for baseline and MHW cruises followed similar (not significantly different) slope relationships to NCD. All biomass components, except Prochlorococcus, were negatively related to NCD, and community biomass structure realigned according to regression slopes differences with NCD variability. Empirically derived biomass‐NCD relationships could be useful for calibrating models that explore future food‐web impacts in this coastal upwelling system. [ABSTRACT FROM AUTHOR]

Biological rate measurements provide critical information for understanding key processes and modeling future states of marine ecosystems. Experimentally derived rates can be challenging to interpret when methodological assumptions are untested or potentially violated under variable natural conditions, such as the assumed linear grazing response of the dilution technique for estimating rates of phytoplankton growth and microzooplankton grazing impact. Here, we show that grazing nonlinearity can be related to the ratio of initial phytoplankton biomass to the half-saturation parameter in the Holling II model, while not being affected by varying grazer biomass during dilution experiments. From this, we present theory to recover growth and grazing rates from multi-treatment dilution experiments with nonlinear grazing results. We test our analyses with data collected during the California Current Ecosystem-Long-Term Ecological Research (CCE-LTER) program. We show that using a linear regression in 2-treatment dilution experiments may lead to underestimates of microzooplankton grazing rates, particularly in high-phytoplankton-biomass coastal regions where grazing can be saturated. Using the Holling II grazing model and a correction factor, growth and grazing rates from 2-treatment experiments can also be estimated, as illustrated by application to Lagrangian water-tracking studies of growth and grazing dynamics in the CCE.

Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg C⋅m-2⋅d-1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional ∼225 mg C⋅m-2⋅d-1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.

Nutrient dynamics, phytoplankton rate processes, and export were examined in a frontal region between an anticyclone and a pair of cyclones 120 km off the coast in the southern California Current System (sCCS). Low silicic acid: nitrate ratios (Si:N) and high nitrate to iron ratios (N:Fe) characteristic of Fe-limiting conditions in the sCCS were associated with the northern cyclone and with the transition zone between the cyclones and the anticyclone. Phytoplankton growth in low-Si:N, high-N:Fe waters responded strongly to added Fe, confirming growth limitation by Fe of the diatom-dominated phytoplankton community. Low Si:N waters had low biogenic silica content, intermediate productivity, but high export compared to intermediate Si:N waters indicating increased export efficiency under Fe stress. Biogenic silica and particulate organic carbon (POC) export were both high beneath low Si:N waters with biogenic silica export being especially enhanced. This suggests that relatively high POC export from low Si:N waters was supported by silica ballasting from Fe-limited diatoms. Higher POC export efficiency in low Si:N waters may have been further enhanced by lower rates of organic carbon remineralization due to reduced grazing of more heavily armored diatoms growing under Fe stress. The results imply that Fe stress can enhance carbon export, despite lowering productivity, by driving higher export efficiency.

In the offshore waters of Southern California, submesoscale processes associated with fronts may stimulate phytoplankton blooms and lead to biomass shifts at multiple trophic levels. Here we report the results of a study on the cycling of biogenic silica (bSiO2) with estimates of the contributions of diatoms to primary and new production in water masses adjacent to (i.e., coastal or oceanic) and within an offshore front in the Southern California Current Ecosystem (CCE). The coastal and oceanic water were sampled in cyclonic and anticyclonic eddies, respectively, with the frontal water being an interaction region between the eddy types. Concentrations of bSiO2 varied by 25-fold across the front, with concentrations in frontal waters 20-25% of those in coastal waters. Rates of biogenic silica production spanned an equally large range, with rates within the frontal region that were half those in the coastal regions. Contributions of diatoms to primary and new production were disproportionately higher than their contribution to autotrophic biomass in all areas, ranging from 5-8%, 19-30%, and 32-43% for both processes in the oceanic, frontal and coastal waters, respectively. Across the frontal area, diatoms could account for

We investigated the spatial variability of picocyanobacterial community structure across the Costa Rica Dome (CRD), an offshore upwelling system characterized by high seasonal abundance of Synechococcus spp. We constructed clone libraries of the rpoC1 gene to survey picocyanobacterial diversity and developed specific real-time quantitative polymerase chain reaction assays to assess the distribution of genetically distinct Synechococcus (SYN) and Prochlorococcus (PRO) populations across vertical and horizontal physicochemical gradients. Flow cytometry data showed that cell abundances for both SYN and PRO were highest near the dome center. Phylogenetic analysis of rpoC1 sequences revealed a remarkably high and distinctive picocyanobacterial diversity (FLU1-3, CRD1, Clade II, XV, XVI) that included "novel" SYN and PRO genotypes. Furthermore, genetically different populations exhibited vertical and horizontal spatial partitioning. Abundances of distinct SYN genotypes peaked at subsequent depth horizons, leading to a fine vertical structure with at least three populations stacked within the upper 30-40 m at the dome. Clade II and FLU1A peaked in surface waters, while maximum concentrations of CRD1, FLU1B, and Clade XVI occurred in the upper and lower thermocline, respectively. Horizontally, Clade II abundance in surface waters remained high across the entire region, while SYN genotypes CRD1 and FLU1A increased with shoaling of the thermo- and nutricline toward the center of the dome to become the dominant genotypes of the SYN assemblage in the dome. Below the mixed layer, Clade XVI and PRO genotype FLU2, virtually absent outside the dome, became abundant components of the picocyanobacterial assemblage. Despite their phylogenetic relatedness, FLU1A and FLU1B subclades followed different distributional patterns, suggesting ecological significance of the microdiversity within the clade. The unprecedented fine vertical structure demonstrated for SYN genotypes is driven by sharp physicochemical gradients (e.g., density, nutrient, oxygen, and trace metals) created by the dome and the presence of a shallow oxycline that enhances habitat diversification. © 2014, by the Association for the Sciences of Limnology and Oceanography, Inc.

A slide preparation method for seawater samples preserved in acid Lugol's is presented here as an alternative to the traditional Utermöhl settling chamber method for microplankton analysis. This preparation maintains the integrity of fragile cells, such as ciliates, resolves issues associated with the transience of samples prepared in settling chambers, and allows the use of automated image acquisition methods. Samples are filtered onto polycarbonate membranes and analyzed with transmitted light microscopy. The visibility of pore outlines is minimized by using mounting oil (Cargille Series A immersion oil, Certified Refractive Index, nD25°C 1.5840 ± 0.0002) with a refractive index matching that of the membrane material. We assessed the efficacy of this new method by comparing abundance and biomass estimates for ciliates in settled and filtered samples. Acceptable results were found for the most delicate of samples stored long-term in acid Lugol's. Some cell shrinkage occurred during the filtration and brief drying steps. Therefore, corrections for ciliate length and width measurements in filtered samples were determined to counteract this effect on total cell biovolume. Overall, the method provides a simple and stable alternative to settling chamber analysis for ciliates preserved in acid Lugol's. © 2014, by the American Society of Limnology and Oceanography, Inc.

In October 2008, we investigated pelagic community composition and biomass, from bacteria to fish, across a sharp frontal gradient overlying deep waters south of Point Conception, California. This northsouth gradient, which we called A-Front, was formed by the eastward flow of the California Current and separated cooler mesotrophic waters of coastal upwelling origin to the north, from warm oligotrophic waters of likely mixed subarcticsubtropical origin to the south. Plankton biomass and phytoplankton growth rates were two to three times greater on the northern side, and primary production rates were elevated 5-fold to the north. Compared with either of the adjacent waters, the frontal interface was strongly enriched and uniquely defined by a subsurface bloom of large diatoms, elevated concentrations of suspension-feeding zooplankton, high bioacoustical estimates of pelagic fish and enhanced bacterial production and phytoplankton biomass and photosynthetic potential. Such habitats, though small in areal extent, may contribute disproportionately and importantly to regional productivity, nutrient cycling, carbon fluxes and trophic ecology. As a general introduction to the A-Front study, we provide an overview of its design and implementation, a brief summary of major findings and a discussion of potential mechanisms of plankton enrichment at the front. © 2012 The Author.

Processes that occur at mesoscale and submesoscale features such as eddies and fronts are important for marine ecosystem dynamics and biogeochemical fluxes. However, their impacts on the fate of biogenic organic carbon in coastal oceans are not well quantified because physical and biological interactions at such features are very complex with short time-and small spatial scales variability. As part of the California Current Ecosystem Long-Term Ecological Research (CCE-LTER) Process studies in the southern California Current in October 2008, we sampled across a strong temperature and chlorophyll front ('A-Front') separating water masses with distinct hydrographic and biogeochemical characteristics and a modified biological assemblage at the frontal interface. Thorpe-scale analyses of the hydrographic data from a free-fall moving vessel profiler suggested an increased diapycnal diffusive nitrate flux at the front zone. Based on these field data, we use data-driven diagnostic biogeochemical models to quantify how the front-induced physical mixing influenced the production, grazing and transport of phytoplankton carbon in the southern California Current. Our results suggest that enhanced diffusive diapycnal fluxes of nutrients stimulated phytoplankton primary production at the front; this effect, together with reduced microzooplankton grazing, increased net growth of the phytoplankton community leading to locally enhanced biomass of large phytoplankton, such as diatoms, in the frontal zone. © 2012 The Author.

Despite the increasing use of linear inverse modeling techniques to elucidate fluxes in undersampled marine ecosystems, the accuracy with which they estimate food web flows has not been resolved. New Markov Chain Monte Carlo (MCMC) solution methods have also called into question the biases of the commonly used L2 minimum norm (L2MN) solution technique. Here, we test the abilities of MCMC and L2MN methods to recover field-measured ecosystem rates that are sequentially excluded from the model input. For data, we use experimental measurements from process cruises of the California Current Ecosystem (CCE-LTER) Program that include rate estimates of phytoplankton and bacterial production, micro- and mesozooplankton grazing, and carbon export from eight study sites varying from rich coastal upwelling to offshore oligotrophic conditions. Both the MCMC and L2MN methods predicted well-constrained rates of protozoan and mesozooplankton grazing with reasonable accuracy, but the MCMC method overestimated primary production. The MCMC method more accurately predicted the poorly constrained rate of vertical carbon export than the L2MN method, which consistently overestimated export. Results involving DOC and bacterial production were equivocal. Overall, when primary production is provided as model input, the MCMC method gives a robust depiction of ecosystem processes. Uncertainty in inverse ecosystem models is large and arises primarily from solution under-determinacy. We thus suggest that experimental programs focusing on food web fluxes expand the range of experimental measurements to include the nature and fate of detrital pools, which play large roles in the model. © 2011 Elsevier B.V.

The high biological production of the California Current System (CCS) results from the seasonal development of equatorward alongshore winds that drive coastal upwelling. While several climatic fluctuation patterns influence the dynamics and biological productivity of the CCS, including the El Niño-Southern Oscillation (ENSO), the Pacific Decadal Oscillation index (PDO) and the North Pacific Gyre Oscillation (NPGO), the mechanisms of interaction between climatic oscillations and the CCS upwelling dynamics have remained obscure. Here, we use Singular Spectral Analysis (SSA) to reveal, for the first time, low-frequency concordance between the time series of climatic indices and upwelling intensity along the coast of western North America. Based on energy distributions in annual, semiannual and low-frequency signals, we can divide the coast into three distinct regions. While the annual upwelling signal dominates the energy spectrum elsewhere, low-frequency variability is maximal in the regions south of 33°N. Non-structured variability associated with storms and turbulent mixing is enhanced at northerly locations. We found that the low-frequency signal is significantly correlated with different climatic indices such as PDO, NPGO and ENSO with the correlation patterns being latitude-dependent. We also analyzed the correlations between this upwelling variability and sea surface temperature (SST) and sea level pressure (SLP) throughout the North Pacific to visualize and interpret the large-scale teleconnection dynamics in the atmosphere that drive the low-frequency coastal winds. These results provide new insights into the underlying mechanisms connecting climatic patterns with upwelling dynamics, which could enhance our prediction and forecast capabilities of the effects of future oceanographic and climatic variability in the CCS.

Experimental studies of phytoplankton growth and grazing processes were conducted in the coastal upwelling system off Point Conception, California to test the hypothesis that phytoplankton growth and grazing losses determine, to first order, the local dynamics of phytoplankton in the upwelling circulation. Eight experiments of 3-5 days each were conducted over the course of two cruises in May-June 2006 and April 2007 following the trajectories of satellite-tracked drifters. Rates of phytoplankton growth and microzooplankton grazing were determined by daily in situ dilution incubations at 8 depths spanning the euphotic zone. Mesozooplankton grazing was assessed by gut fluorescence analysis of animals collected from net tows through the euphotic zone. We compared directly the net rates of change observed for the ambient phytoplankton community to the net growth rates predicted from experimental determinations of each process rate. The resulting relationship accounted for 91% of the variability observed, providing strong support for the growth-grazing hypothesis. In addition, grazing by mesozooplankton was unexpectedly high and variable, driving a substantial positive to negative shift in phytoplankton net rate of change between years despite comparable environmental conditions and similar high growth rates and suggesting strong top-down control potential. The demonstrated agreement between net ambient and experimental community changes is an important point of validation for using field data to parameterize models. Data sets of this type may provide an important source of new information and rate constraints for developing better coupled biological-physical models of upwelling system dynamics. © 2009 Elsevier Ltd. All rights reserved.

In this paper, we review the state of the art and major challenges in current efforts to incorporate biogeochemical functional groups into models that can be applied on basin-wide and global scales, with an emphasis on models that might ultimately be used to predict how biogeochemical cycles in the ocean will respond to global warming. We define the term "biogeochemical functional group" to refer to groups of organisms that mediate specific chemical reactions in the ocean. Thus, according to this definition, "functional groups" have no phylogenetic meaning-these are composed of many different species with common biogeochemical functions. Substantial progress has been made in the last decade toward quantifying the rates of these various functions and understanding the factors that control them. For some of these groups, we have developed fairly sophisticated models that incorporate this understanding, e.g. for diazotrophs (e.g. Trichodesmium), silica producers (diatoms) and calcifiers (e.g. coccolithophorids and specifically Emiliania huxleyi). However, current representations of nitrogen fixation and calcification are incomplete, i.e., based primarily upon models of Trichodesmium and E. huxleyi, respectively, and many important functional groups have not yet been considered in open-ocean biogeochemical models. Progress has been made over the last decade in efforts to simulate dimethylsulfide (DMS) production and cycling (i.e., by dinoflagellates and prymnesiophytes) and denitrification, but these efforts are still in their infancy, and many significant problems remain. One obvious gap is that virtually all functional group modeling efforts have focused on autotrophic microbes, while higher trophic levels have been completely ignored. It appears that in some cases (e.g., calcification), incorporating higher trophic levels may be essential not only for representing a particular biogeochemical reaction, but also for modeling export. Another serious problem is our tendency to model the organisms for which we have the most validation data (e.g., E. huxleyi and Trichodesmium) even when they may represent only a fraction of the biogeochemical functional group we are trying to represent. When we step back and look at the paleo-oceanographic record, it suggests that oxygen concentrations have played a central role in the evolution and emergence of many of the key functional groups that influence biogeochemical cycles in the present-day ocean. However, more subtle effects are likely to be important over the next century like changes in silicate supply or turbulence that can influence the relative success of diatoms versus dinoflagellates, coccolithophorids and diazotrophs. In general, inferences drawn from the paleo-oceanographic record and theoretical work suggest that global warming will tend to favor the latter because it will give rise to increased stratification. However, decreases in pH and Fe supply could adversely impact coccolithophorids and diazotrophs in the future. It may be necessary to include explicit dynamic representations of nitrogen fixation, denitrification, silicification and calcification in our models if our goal is predicting the oceanic carbon cycle in the future, because these processes appear to play a very significant role in the carbon cycle of the present-day ocean and they are sensitive to climate change. Observations and models suggest that it may also be necessary to include the DMS cycle to predict future climate, though the effects are still highly uncertain. We have learned a tremendous amount about the distributions and biogeochemical impact of bacteria in the ocean in recent years, yet this improved understanding has not yet been incorporated into many of our models. All of these considerations lead us toward the development of increasingly complex models. However, recent quantitative model intercomparison studies suggest that continuing to add complexity and more functional groups to our ecosystem models may lead to decreases in predictive ability if the models are not properly constrained with available data. We also caution that capturing the present-day variability tells us little about how well a particular model can predict the future. If our goal is to develop models that can be used to predict how the oceans will respond to global warming, then we need to make more rigorous assessments of predictive skill using the available data. © 2006 Elsevier Ltd. All rights reserved.