The coastal upwelling ecosystems analysis program as an experience in international cooperation

Abstract

The Coastal Upwelling Ecosystems Analysis CUEA) Project, the major effort in the Living Resources part of the United States Program for the International Decade of Ocean Exploration (IDOE), included a variety of approaches to international involvement. Escalating cost, expanded geographical scope, and increasing insistence on coastal state participation in research projects off foreign shores heighten the likelihood that similar large‐scale, long‐term research efforts will require particular attention to international cooperation and collaboration. A look at the international aspects of CUEA is timely and instructive. This discussion reviews the background to the upwelling project, the nature of international involvement, and closes with some observations about the requirements for similar projects in the future.     

Diel and daily fluctuations in virioplankton production in coastal ecosystems

Viruses saturate the world around us, yet a basic understanding of how viral impacts on microbial host organisms vary over days to hours, which typify the replication cycles of aquatic viruses, remains elusive. Thus, diel patterns of viral production (VP) in Chesapeake Bay surface waters were examined on five sampling dates. Day-to-day variations in VP in the Chesapeake and coastal California surface waters were also investigated. Significant variations in VP were detected over 24 h cycles during four of five studies, but rates did not vary significantly over the course of a few days in either location. Diel patterns of VP displayed seasonality with shorter viral assemblage turnover times and shorter times to maximum viral abundance in summer, implying shorter replication cycles for virus–host systems in warmer months. No correlation was found between VP and time of day, likely due to seasonal changes in the diel patterns of VP. This analysis significantly increases our knowledge of the short-term patterning of in situ VP, and thus viral impacts, and suggests that variations in viral biology in response to changes in host communities or physio-chemical properties affect both diel and seasonal cycles and magnitudes of VP.     

Greenhouse gas,upwelling‐favorable winds,and the future of coastal ocean upwelling ecosystems

Coastal ocean upwelling ecosystems generally represent the most productive large marine ecosystems of the world's oceans, in terms of both primary production rates and tonnages of exploitable fish produced. The Peruvian upwelling system, in particular, stands out as a major factor in world fish production. The Pacific trade winds have traditionally been considered to be the primary driving force for the upwelling system off Peru, but are projected to weaken as climate change proceeds. This leads to concern that the upwelling process in the Peru system, to which its productivity is linked, may likewise weaken. However, other mechanisms involving greenhouse‐associated intensification of thermal low‐pressure cells over the coastal landmasses of upwelling regions suggest general intensification of wind‐driven ocean upwelling in coastal upwelling regions of the world's oceans. But although certain empirical results have supported this expectation, it has not been consistently corroborated in climate model simulations, possibly because the scale of the coastal intensification may be small relative to the scales that are appropriately reflected in the standard models. Here we summarize available evidence for the intensification mechanism and present a proxy test that uses variations in water vapor, the dominant natural greenhouse gas, to offer multiple‐realization empirical evidence for action of the proposed mechanism in the real world situation. While many potential consequences to the future of marine ecosystems would codepend on climate change‐related changes in the thermocline and nutricline structures, an important subset, involving potential increased propensities for hypoxia, noxious gas eruptions, toxic red tide blooms, and/or jellyfish outbreaks, may depend more directly on changes in the upwelling‐favorable wind itself. A prospective role of fisheries in either mitigating or reinforcing this particular class of effects is suggested.     

Seasonality in coastal benthic ecosystems

For historical reasons, knowledge about seasonality in the dynamics of marine benthic suspension feeders from temperate areas comes mainly from studies of cold temperate seas. Recent surveys of Mediterranean taxa show different patterns from those observed in cold temperate seas, which are characterized by winter dormancy. In the Mediterranean, summer dormancy predominates among taxa and appears to be related to energetic constraints. Temperature and food availability are crucial to the dynamics of benthic suspension feeders. However, because these factors tend to be positively correlated in cold temperate seas, it is difficult to distinguish between their effects. Such correlation does not occur in Mediterranean ecosystems. The contrast between recent studies in the Mediterranean and in other areas can help to disentangle confounded environmental controls.     

Sulfate reduction in coastal ecosystems

The diversity and activity of dissimilatory Fe(III)-reducing bacteria was investigated in acidic, ochre-precipitating springs on Mam Tor, East Midlands, UK. The springs at this acid rock drainage site are located below a 3000 year old landslip, where biooxidation of exposed pyrite-containing minerals has resulted in the production of metal-laden acidic waters. A diverse microbial community was found downstream in the sediments dominated by Fe(III) minerals, and included close relatives to known acidophilic (Acidimicrobium and Acidiphilium) and neutraphilic (Geobacter and Pelobacter) Fe(III)-reducing bacteria. Analysis by XRD and TEM confirmed the presence of both amorphous and well-defined Fe(III) mineral phases in the sediments including lepidocrocite, goethite and schwertmannite. Microcosm-based experiments demonstrated that the bioavailable Fe(III) was reduced under anaerobic conditions, concomitant with sulphate release. XRD analysis suggested that schwertmannite (an iron sulphate hydroxide) was utilized preferentially by the Fe(III)-reducing bacteria, leading to the release of sulphate. Although the microcosms contained sufficient concentrations of naturally occurring electron donor to sustain significant levels of Fe(III) reduction, this process was stimulated by the addition of glycerol and complex electron donors. Thus, the acidic Fe(III)-containing sediments contain a diversity of DIRBs that can be stimulated by the addition of electron donor as a first step in the reversal of acid rock and acid mine drainage contamination.     

Desertification alters patterns of aboveground net primary production in Chihuahuan ecosystems

Abstract The Chihuahuan desert of New Mexico, USA, has changed in historical times from semiarid grassland to desert shrublands dominated by Larrea tridentata and Prosopis glandulosa. Similar displacement of perennial grasslands by shrubs typifies desertification in many regions. Such structural vegetation change could alter average values of net primary productivity, as well as spatial and temporal patterns of production. We investigated patterns of aboveground plant biomass and net primary production in five ecosystem types of the Jornada Basin Long‐Term Ecological Research (LTER) site. Comparisons of shrub‐dominated desertified systems and remnant grass‐dominated systems allowed us to test the prediction that shrublands are more heterogeneous spatially, but less variable over time, than grasslands. We measured aboveground plant biomass and aboveground net primary productivity (ANPP) by species, three times per year for 10 years, in 15 sites of five ecosystem types (three each in Larrea shrubland, Bouteloua eriopoda grassland, Prosopis dune systems, Flourensia cernua alluvial flats, and grass‐dominated dry lakes or playas). Spatial heterogeneity of biomass at the scale of our measurements was significantly greater in shrub‐dominated systems than in grass‐dominated vegetation. ANPP was homogeneous across space in grass‐dominated systems, and in most growing seasons was significantly more patchy in shrub vegetation. Substantial interannual variability in ANPP complicates comparison of mean values across ecosystem types, but grasslands tended to support higher ANPP values than did shrub‐dominated systems. There were significant interactions between ecosystem type and season. Grasslands demonstrated higher interannual variation than did shrub systems. Desertification has apparently altered the seasonality of productivity in these systems; grasslands were dominated by summer growth, while sites dominated by Larrea or Prosopis tended to have higher spring ANPP. Production was frequently uncorrelated across sites of an ecosystem type, suggesting that factors other than season, regional climate, or dominant vegetation may be significant determinants of actual NPP.     

Damped trophic cascades driven by fishing in model marine ecosystems

The largest perturbation on upper trophic levels of many marine ecosystems stems from fishing. The reaction of the ecosystem goes beyond the trophic levels directly targeted by the fishery. This reaction has been described either as a change in slope of the overall size spectrum or as a trophic cascade triggered by the removal of top predators. Here we use a novel size- and trait-based model to explore how marine ecosystems might react to perturbations from different types of fishing pressure. The model explicitly resolves the whole life history of fish, from larvae to adults. The results show that fishing does not change the overall slope of the size spectrum, but depletes the largest individuals and induces trophic cascades. A trophic cascade can propagate both up and down in trophic levels driven by a combination of changes in predation mortality and food limitation. The cascade is damped as it comes further away from the perturbed trophic level. Fishing on several trophic levels leads to a disappearance of the signature of the trophic cascade. Differences in fishing patterns among ecosystems might influence whether a trophic cascade is observed.     

Nitrogen cycling in coastal marine ecosystems

It is generally considered that nitrogen availability is one of the major factors regulating primary production in temperate coastal marine environments. Coastal regions often receive large anthropogenic inputs of nitrogen that cause eutrophication. The impact of these nitrogen additions has a profound effect in estuaries and coastal lagoons where water exchange is limited. Such increased nutrient loading promotes the growth of phytoplankton and fast growing pelagic macroalgae while rooted plants (sea-grasses) and benthic are suppressed due to reduced light availability. This shift from benthic to pelagic primary production introduces large diurnal variations in oxygen concentrations in the water column. In addition oxygen consumption in the surface sediments increases due to the deposition of readily degradable biomass. In this review the physico-chemical and biological factors regulating nitrogen cycling in coastal marine ecosystems are considered in relation to developing effective management programmes to rehabilitate seagrass communities in lagoons currently dominated by pelagic macroalgae and/or cyanobacteria.     

Bivalve carrying capacity in coastal ecosystems

The carrying capacity of suspension feeding bivalvesin 11 coastal and estuarine ecosystems is examined. Bivalve carrying capacity is defined in terms of watermass residence time, primary production time andbivalve clearance time. Turnover times for the 11ecosystems are compared both two and threedimensionally. Fast systems, e.g., Sylt and NorthInlet, have turnover times of days or less, while,slow systems, e.g., Delaware Bay, have turnover timesof months and years. Some systems,Marennes-Oléron, South San Francisco Bay and NorthInlet, require a net influx of phytoplankton in orderto support their bivalve populations. Three systems,Carlingford Lough, Chesapeake Bay and Delaware Bay,have very long bivalve clearance times due to small orreduced bivalve filter feeder populations. Carlingford Lough stands out because it is a naturallyplanktonic system now being converted to bivalveculture with an adherently stronger benthic-pelagiccoupling. Existing models of bivalve carrying capacity arereviewed. The Herman model is utilized as anappropriate ecosystem level model to examine carryingcapacity because it includes the three major turnovertime elements of water mass residence time, primaryproduction time and bivalve filter feeder clearancetime. The graphical analysis suggests that massive andsuccessful bivalve filter feeder populations are foundin systems with relatively short residence times(<40 days) and short primary production times (<4days) in order to sustain a high bivalve biomass withits associated rapid clearance times. Outliersystems are constrained by long water mass residencetimes, extended primary production times, and longclearance times. This revised version was published online in August 2006 with corrections to the Cover Date.     

Progress in management of coastal ecosystems

The process of coastal zone management in the United States has progressed to the point that it is now possible to embrace whole coastal ecosystems in management programs. The structural and dynamical features of the ecosystems have to be known before management can succeed. Furthermore, it is necessary to disaggregate the systems into subsystems. The following six subsystems have been used as the best compromise between scientific and administrative needs: the watershed terrain, the land drainage system, the coastal basin, the basin floor, the coastal waters, and the ocean. The Apalachicola National Estuarine Sanctuary in Florida is used as a case history of a managed ecosystem.     

Changes in phytoplankton ecophysiology across a coastal upwelling front

The abundance, taxonomic composition and patterns of macromolecularsynthesis of phytoplankton were determined across an upwelling-inducedthermal front in the central Cantabrian Sea (southern Bay ofBiscay) during July 1993. Enhanced levels of phytoplankton biomass,diatom abundance and photosynthetic rate were measured on thecoastal side of the front. Relative carbon (C) incorporationinto proteins increased noticeably on the oceanic side, takingvalues of up to 64%, whereas changes in the relative C incorporationinto lipids and low-molecular-weight metabolites followed anopposite trend. Phytoplankton cells on the oceanic side of thefront were adapted to the prevailing growth-limiting conditionsby maintaining the synthesis of functionally essential molecules—proteins—ratherthan the synthesis of storage compounds. As a result, the carbonto nitrogen uptake ratio varied from {small tilde}5.7 in offshorewaters to 8.0 in the nearshore region. Our results suggest thatthe taxonomic and physiological changes in phytoplankton assemblagesas a response to upwelling may result in an increase in thesynthesis of organic C relative to the upward flux of nitrate.     

Phytoplankton biomass and production in shelf waters off NW Spain: spatial and seasonal variability in relation to upwelling

Summary Chlorophyll-a and primary production on the euphotic zone of the N-NW Spanish shelf were studied at 125 stations between 1984 and 1992. Three geographic areas (Cantabrian Sea, Rías Altas and Was Baixas), three bathymetric ranges (20 to 60 m, 60 to 150 m and stations deeper than 200 m), and four oceanographic stages (spring and autumn blooms, summer upwelling, summer stratification and winter mixing) were considered. One of the major sources of variability of chlorophyll and production data was season. Bloom and summer upwelling stages have equivalent mean and maximum values. Average chlorophyll-a concentrations approximately doubled in every step of the increasing productivity sequence: winter mixing — summer stratification — high productivity (upwelling and bloom) stages. Average primary production rates increased only 60% in the described sequence. Mean (± sd) values of chlorophyll-a and primary production rates during the high productivity stages were 59.7 ± 39.5 mg Chl-a m^–2 and 86.9 ± 44.0 mg C m^–2 h^–1, respectively. Significant differences in both chlorophyll and primary production resulted between geographic areas in most stages. Only 27 stations showed the effects of the summer upwelling that affected coastal areas in the Cantabrian Sea and Rías Baixas shelf, but also shelf-break stations in the Rías Altas area. The Rías Baixas area had lower chlorophyll than both the Rías Altas and the Cantabrian Sea areas during spring and autumn blooms, but higher during summer upwelling events. On the contrary, primary production rates were higher in the Rías Baixas area during blooms in spring and autumn. Mid-shelf areas showed the highest chlorophyll concentrations during high productivity stages, probably due to the existence of frontal zones in all geographic areas considered. The estimated phytoplankton growth rates were comparable to those of other coastal upwelling systems, with average values lower than the maximum potential growth rates. Doubling rates for upwelling and stratification stages in the northern and Rías Altas shelf areas were equivalent, despite larger biomass accumulations during upwelling events. Low turnover rates of the existing biomass in the Rías Baixas shelf in upwelling stages suggests that the accumulation of phytoplankton was due mainly to the export from the highly productive rías, while the contribution of in situ production to these accumulations was relatively lower.     

Consumption of aquatic subsidies by soil invertebrates in coastal ecosystems

Routes of aquatic allochthonous inputs (aquatic subsidies) to detrital food webs are studied, as is the effect of aquatic subsidies on the functional and taxonomic structure of soil invertebrate communities in coastal ecosystems. The study took place in the coastal zone of an oxbow lake of the Pra River in the Oka Reserve. The results indicate a strong dependence of soil animals in the coastal habitats on aquatic subsidies. Isotopic analysis shows that aquatic resources enter soil food webs not only via predators feeding on flying insects or aquatic prey, but also via saprophages decomposing organic debris of aquatic origin. The contribution of aquatic subsidies to the energy balance of soil invertebrates decreases rapidly with increasing distance from the lake. The fraction of aquatic carbon in tissues of collembolans and saprophages is negligible already a few meters from the water edge. The dependence of predatory invertebrates on aquatic resources can be traced at somewhat greater distance (tens of meters).     

Climatically driven fluctuations in Southern Ocean ecosystems

Determining how climate fluctuations affect ocean ecosystems requires an understanding of how biological and physical processes interact across a wide range of scales. Here we examine the role of physical and biological processes in generating fluctuations in the ecosystem around South Georgia in the South Atlantic sector of the Southern Ocean. Anomalies in sea surface temperature (SST) in the South Pacific sector of the Southern Ocean have previously been shown to be generated through atmospheric teleconnections with El Niño Southern Oscillation (ENSO)-related processes. These SST anomalies are propagated via the Antarctic Circumpolar Current into the South Atlantic (on time scales of more than 1 year), where ENSO and Southern Annular Mode-related atmospheric processes have a direct influence on short (less than six months) time scales. We find that across the South Atlantic sector, these changes in SST, and related fluctuations in winter sea ice extent, affect the recruitment and dispersal of Antarctic krill. This oceanographically driven variation in krill population dynamics and abundance in turn affects the breeding success of seabird and marine mammal predators that depend on krill as food. Such propagating anomalies, mediated through physical and trophic interactions, are likely to be an important component of variation in ocean ecosystems and affect responses to longer term change. Population models derived on the basis of these oceanic fluctuations indicate that plausible rates of regional warming of 1^oC over the next 100 years could lead to more than a 95% reduction in the biomass and abundance of krill across the Scotia Sea by the end of the century.     

Ecological drivers of plant diversity patterns in remnants coastal sand dune ecosystems along the northern Adriatic coastline

Coastal sand dunes represent one of the most fragile ecosystems in the Mediterranean basin. These habitats naturally suffer the action of several limiting factors such as sand burial, marine aerosol and low soil fertility; on the other hand, they often host species of high conservation value. Over the last decades, they have also experienced a high level of biological invasion. In this study, we sampled psammophilous vegetation in two sites in the northern Adriatic coast belonging to the Natura 2000 network to describe diversity patterns and to identify the main ecological drivers of species diversity. Plant species richness and their abundance were assessed in each plot. Differences in species composition for native and alien species were compared via PERMANOVA analysis. Species complementarity was explored by partitioning beta diversity in its spatial components (richness and replacement). A Generalized Linear Model was also computed to assess the main environmental factors that may promote invasiveness in these ecosystems. For the investigated area, our results highlight the strong differentiation in community composition both in alien and native species: in particular alien species showed on average a lower complementarity among habitats compared to native species. Specifically, communities seem to be more diversified when larger spatial scales were considered. Beta diversity in both groups appears to be more dominated by the richness component with respect to the replacement component. Furthermore, in these habitats, the occurrence of alien species was shown to be related to geomorphological predictors more than climatic variables.     

Diatom taphocoenoses in the coastal upwelling area off South West Africa

The diatom floral composition of 124 sediment samples from the South East Atlantic records the influence of coastal upwelling on sediment composition off South West Africa. Inner shelf samples between 19° and 24°S are rich in diatom valves and the patterns of diatom species distribution in these samples are related to the coastal upwelling process. Comparison with recent phytoplankton data shows that the sediment assemblages preserve many of the important species of the diatom biocoenoses, including Chaetoceros (resting spores), Delphineis karstenii, Thalassiosira eccentrica, and Thalassionema nitzschioides. Delphineis karstenii, a pioneer species in enriched coastal water, occurs nearshore and Chaetoreros resting spores are widespread, with highest relative abundance values in some more offshore samples. The abundance of the Thalassiosira eccentrica group and of Thalassionema nitzschioides in sediment samples in and near Walvis Bay reflects the recurrence of intense upwelling off this part of the coast. An abundance of large centric species has been reported in hydrological conditions characteristic of newly upwelled waters and, correspondingly, Actinocyclus octonarius and some large Coscinodiscus species occur in the sediments in nearshore patches or belts.     

Global patterns of root turnover for terrestrial ecosystems

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands ( r ² = 0.48) and forests ( r ² = 0.17) and for total root biomass in shrublands ( r ² = 0.55). On the basis of the best-fit exponential model, the Q ₁₀ for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature ( r ² = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.     

The ups and downs of trophic control in continental shelf ecosystems

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.