Spatial Structure of Pelagic Ecosystem Processes in the Global Ocean

We propose an operational definition of spatial structure in the oceanic ecosystem; it is equivalent to the large-scale, horizontal distribution of the dominant ecophysiological rate parameters for the questions of interest. In an important, particular case, the relevant rate parameters are those that characterize autotrophic production. In the ocean, these parameters are believed to be distributed in a manner that is not smoothly continuous. Rather, they seem to have a piecewise continuous distribution. This leads to a requirement to partition the ocean into a suite of provinces where the boundaries between the provinces mark the locations of abrupt changes in the magnitudes of the rate parameters. The area covered by a particular province represents an area of common physical forcing, insofar as the forcing is relevant to autotrophic production. The boundaries are taken to be elastic rather than fixed, such that they can respond to variations in forcing. At any given time, the boundaries can be located with the aid of remotely sensed imagery, especially ocean-color imagery. Received 7 January 1999; accepted 2 June 1999.     

The Role of Sharks and Longline Fisheries in a Pelagic Ecosystem of the Central Pacific

The increased exploitation of pelagic sharks by longline fisheries raised questions about changes in the food webs that include sharks as apex predators. We used a version of Ecopath/Ecosim models to evaluate changes in trophic interactions due to shark exploitation in the Central North Pacific. Fisheries targeted on blue sharks tend to produce compensatory responses that favor other shark species and billfishes, but they have only modest effects on the majority of food web components. Modest levels of intraguild predation (adult sharks that eat juvenile sharks) produce strong, nonlinear responses in shark populations. In general, analysis of the Central North Pacific model reveals that sharks are not keystone predators, but that increases in longline fisheries can have profound effects on the food webs that support sharks. Received 19 April 2001; accepted 2 October 2001.     

Selenium and Mercury in Pelagic Fish in the Central North Pacific Near Hawaii

Protective effects of selenium against mercury toxicity have been demonstrated in all animal models evaluated. As interactions between selenium and mercury and their molar ratios in seafood are essential factors in evaluating risks associated with dietary mercury exposure, considering mercury content alone is inadequate. In this study, the absolute and molar concentrations of mercury and selenium were determined in edible portions from 420 individual fish representing 15 species of pelagic fish collected from the central North Pacific Ocean near Hawaii. Selenium was in molar excess of mercury in almost all fish species evaluated. The rank order of mean Se/Hg molar ratios was striped marlin (17.6) > yellowfin tuna (14.1) > mahimahi (13.1) > skipjack tuna (12.8) > spearfish (11.4) > wahoo (10.8) > sickle pomfret (6.7) > albacore tuna (5.3) > bigeye tuna (5.2) > blue marlin (4.1) > escolar (2.4) > opah (2.3) > thresher shark (1.5) > swordfish (1.2) > mako shark (0.5). With a Se/Hg molar ratio of less than 1, mako shark was the only fish containing a net molar excess of mercury. A selenium health benefit value based on the absolute amounts and relative proportions of selenium and mercury in seafood is proposed as a more comprehensive seafood safety criterion.     

Pigmented Nanoflagellates Grazing on <Emphasis Type="Italic">Synechococcus</Emphasis>: Seasonal Variations and Effect of Flagellate Size in the Coastal Ecosystem of Subtropical Western Pacific

We investigated seasonal variation of grazing impact of the pigmented nanoflagellates (PNF) with different sizes upon Synechococcus in the subtropical western Pacific coastal waters using grazing experiments with fluorescently labeled Synechococcus (FLS). For total PNF, conspicuous seasonal variations of ingestion rates on Synechococcus were found, and a functional response was observed. To further investigate the impact of different size groups, we separated the PNF into four categories (<3, 3–5, 5–10, and >10 μm). Our results indicated that the smallest PNF (<3 μm PNF) did not ingest FLS and was considered autotrophic. PNF of 3–5 μm in size made up most of the PNF community; however, their ingestion on Synechococcus was too low (0.1–1.9 Syn PNF⁻¹ h⁻¹) to support their growth, and they had to depend on other prey or photosynthesis to survive. The ingestion rate of the 3–5 μm group exhibited no significant seasonal variation; by contrast, the ingestion rates of 5–10 and >10 μm PNFs showed significant seasonal variation. During the warm season, 3–5 μm PNF were responsible for the grazing of 12% of Synechococcus production, 5–10 μm PNF for 48%, and >10 μm PNF for 2%. Taken together, our results demonstrate that the PNF of 3–10 μm consumed most Synechococcus during the warm season and exhibited a significant functional response to the increase in prey concentration.     

Spatial and Temporal Variability in the Ecosystem Metabolism of a High-elevation Lake: Integrating Benthic and Pelagic Habitats

We characterized spatial and temporal variability in net ecosystem production (NEP), community respiration (CR), and gross primary production (GPP) over an ice-free season in an oligotrophic high-elevation lake using high-frequency measurements of dissolved oxygen. We combined the use of free-water and incubation chamber measurements to compare pelagic and benthic habitats and estimate their relative contributions to whole-lake metabolism. Despite a brief period of predominant heterotrophy after snowmelt, both free-water and incubation chamber measurements confirmed autotrophy of the epilimnion in all habitats throughout the ice-free season. In contrast, benthic incubation chambers showed the benthos to be consistently heterotrophic. Although temperature was the strongest seasonal driver of benthic metabolism, bacterioplankton density and indexes of organic matter quality explained the most variability in pelagic metabolism. Driven largely by benthic metabolism, free-water measurements of GPP and CR were twice as high in littoral than pelagic habitats. However, rates of water column primary production overlying the littoral benthos were high enough to overcome net benthic heterotrophy, and seasonal mean NEP in littoral habitats remained positive and not significantly different from pelagic habitats. Benthic rates averaged about 25% of whole-lake metabolism. Pelagic metabolism measurements were affected by littoral rates about half the time, with the degree of isolation between the two a function of advection and water column stability. These results emphasize the importance of characterizing spatial and temporal variability in metabolism within the context of physical dynamics and challenge the notion that benthic metabolism will necessarily be larger than pelagic metabolism in oligotrophic lakes.     

Genetic Diversity and Population Structure of the Pelagic Thresher Shark (Alopias pelagicus) in the Pacific Ocean: Evidence for Two Evolutionarily Significant Units

There has been an increasing concern about shark overexploitation in the last decade, especially for open ocean shark species, where there is a paucity of data about their life histories and population dynamics. Little is known regarding the population structure of the pelagic thresher shark, Alopias pelagicus. Though an earlier study using mtDNA control region data, showed evidence for differences between eastern and western Pacific populations, the study was hampered by low sample size and sparse geographic coverage, particularly a lack of samples from the central Pacific. Here, we present the population structure of Alopias pelagicus analyzing 351 samples from six different locations across the Pacific Ocean. Using data from mitochondrial DNA COI sequences and seven microsatellite loci we found evidence of strong population differentiation between western and eastern Pacific populations and evidence for reciprocally monophyly for organelle haplotypes and significant divergence of allele frequencies at nuclear loci, suggesting the existence of two Evolutionarily Significant Units (ESU) in the Pacific Ocean. Interestingly, the population in Hawaii appears to be composed of both ESUs in what seems to be clear sympatry with reproductive isolation. These results may indicate the existence of a new cryptic species in the Pacific Ocean. The presence of these distinct ESUs highlights the need for revised management plans for this highly exploited shark throughout its range.     

Burning Fire-Prone Mediterranean Shrublands: Immediate Changes in Soil Microbial Community Structure and Ecosystem Functions

Wildfires subject soil microbes to extreme temperatures and modify their physical and chemical habitat. This might immediately alter their community structure and ecosystem functions. We burned a fire-prone shrubland under controlled conditions to investigate (1) the fire-induced changes in the community structure of soil archaea, bacteria and fungi by analysing 16S or 18S rRNA gene amplicons separated through denaturing gradient gel electrophoresis; (2) the physical and chemical variables determining the immediate shifts in the microbial community structure; and (3) the microbial drivers of the change in ecosystem functions related to biogeochemical cycling. Prokaryotes and eukaryotes were structured by the local environment in pre-fire soils. Fire caused a significant shift in the microbial community structure, biomass C, respiration and soil hydrolases. One-day changes in bacterial and fungal community structure correlated to the rise in total organic C and NO(3)(-)-N caused by the combustion of plant residues. In the following week, bacterial communities shifted further forced by desiccation and increasing concentrations of macronutrients. Shifts in archaeal community structure were unrelated to any of the 18 environmental variables measured. Fire-induced changes in the community structure of bacteria, rather than archaea or fungi, were correlated to the enhanced microbial biomass, CO(2) production and hydrolysis of C and P organics. This is the first report on the combined effects of fire on the three biological domains in soils. We concluded that immediately after fire the biogeochemical cycling in Mediterranean shrublands becomes less conservative through the increased microbial biomass, activity and changes in the bacterial community structure.     

Biotic and Abiotic Changes in Ecosystem Structure over a Shrub-Encroachment Gradient in the Southwestern USA

In this study, we investigate changes in ecosystem structure that occur over a gradient of land-degradation in the southwestern USA, where shrubs are encroaching into native grassland. We evaluate a conceptual model which posits that the development of biotic and abiotic structural connectivity is due to ecogeomorphic feedbacks. Three hypotheses are evaluated: 1. Over the shrub-encroachment gradient, the difference in soil properties under each surface-cover type will change non-linearly, becoming increasingly different; 2. There will be a reduction in vegetation cover and an increase in vegetation-patch size that is concurrent with an increase in the spatial heterogeneity of soil properties over the shrub-encroachment gradient; and 3. Over the shrub-encroachment gradient, the range at which soil properties are autocorrelated will progressively exceed the range at which vegetation is autocorrelated. Field-based monitoring of vegetation and soil properties was carried out over a shrub-encroachment gradient at the Sevilleta National Wildlife Refuge in New Mexico, USA. Results of this study show that vegetation cover decreases over the shrub-encroachment gradient, but vegetation-patch size increases, with a concurrent increase in the spatial heterogeneity of soil properties. Typically, there are significant differences in soil properties between non-vegetated and vegetated surfaces, but for grass and shrub patches, there are only significant differences for the biotic soil properties. Results suggest that it is the development of larger, well-connected, non-vegetated patches that is most important in driving the overall behavior of shrub-dominated sites. Results of this study support the hypothesis that feedbacks of functional connectivity reinforce the development of structural connectivity, which increases the resilience of the shrub-dominated state, and thus makes it harder for grasses to re-establish and reverse the vegetation change.     

Structural Compositions of Pelagic Communities in the River Warnow and their Changes

The phytoplankton and heterotrophic microbial communities from the Oberwarnow and Unterwarnow were analyzed from November 1987 to May 1988. The production potential of the microorganisms was quantified in laboratory experiments. On the average, phytoplankton accounted for 73 % of the total biomass in the Oberwarnow and 59 % in the Unterwarnow. The corresponding figures for bacterial biomass were 24 % and 33 %, respectively. If the nutrient supply is unlimited and phytoplankton biomass predominates, the development of primary producers in eutrophic to polytrophic waters is not regulated by metazooplankton. Top-down regulation is possible only if prevailing conditions limit the growth of the dominant phytoplankton species.     

Modelling the vertical distribution of Prochlorococcus and Synechococcus in the North Pacific Subtropical Ocean

A simple model was developed to examine the vertical distribution of Prochlorococcus and Synechococcus ecotypes in the water column, based on their adaptation to light intensity. Model simulations were compared with a 14-year time series of Prochlorococcus and Synechococcus cell abundances at Station ALOHA in the North Pacific Subtropical Gyre. Data were analysed to examine spatial and temporal patterns in abundances and their ranges of variability in the euphotic zone, the surface mixed layer and the layer in the euphotic zone but below the base of the mixed layer. Model simulations show that the apparent occupation of the whole euphotic zone by a genus can be the result of a co-occurrence of different ecotypes that segregate vertically. The segregation of ecotypes can result simply from differences in light response. A sensitivity analysis of the model, performed on the parameter alpha (initial slope of the light-response curve) and the DIN concentration in the upper water column, demonstrates that the model successfully reproduces the observed range of vertical distributions. Results support the idea that intermittent mixing events may have important ecological and geochemical impacts on the phytoplankton community at Station ALOHA.     

Spatial Structure and Distribution of Small Pelagic Fish in the Northwestern Mediterranean Sea

Understanding the ecological and anthropogenic drivers of population dynamics requires detailed studies on habitat selection and spatial distribution. Although small pelagic fish aggregate in large shoals and usually exhibit important spatial structure, their dynamics in time and space remain unpredictable and challenging. In the Gulf of Lions (north-western Mediterranean), sardine and anchovy biomasses have declined over the past 5 years causing an important fishery crisis while sprat abundance rose. Applying geostatistical tools on scientific acoustic surveys conducted in the Gulf of Lions, we investigated anchovy, sardine and sprat spatial distributions and structures over 10 years. Our results show that sardines and sprats were more coastal than anchovies. The spatial structure of the three species was fairly stable over time according to variogram outputs, while year-to-year variations in kriged maps highlighted substantial changes in their location. Support for the McCall''s basin hypothesis (covariation of both population density and presence area with biomass) was found only in sprats, the most variable of the three species. An innovative method to investigate species collocation at different scales revealed that globally the three species strongly overlap. Although species often co-occurred in terms of presence/absence, their biomass density differed at local scale, suggesting potential interspecific avoidance or different sensitivity to local environmental characteristics. Persistent favourable areas were finally detected, but their environmental characteristics remain to be determined.     

Comparison of the Benthic Size Structure Between Sublittoral,Upper-slope and Deep-Sea Areas of the Western Pacific

Benthic size structures were compared between sublittoral, upper-slope and deep-sea areas of the Western Pacific using the mesh-size fractionated distribution of organisms. The average size of the individuals became smaller with increasing water depth, suggesting that the dwarfism is a more common phenomenon than the gigantism in the deep sea. Nonetheless, the mesh size of 0.037 mm was fine enough to retain meiobenthic specimens quantitatively. When the whole benthos was divided into macro- and meiofauna, the size distributions overlapped considerably in a mesh-size fraction between 1.0 and 0.5 mm. In this fraction, the meiofauna predominated over macrofauna in the deep sea, while the opposite applied to the shallower depths. From these results, recommended mesh sizes of the sieves for the studies of macro- and meiobenthos were proposed.     

Historical Changes in Marine Resources,Food-web Structure and Ecosystem Functioning in the Adriatic Sea,Mediterranean

The Mediterranean Sea has been strongly influenced by human activities for millennia. Although the environmental history of its surrounding terrestrial ecosystems has received considerable study, historical changes in its marine realm are less known. We used a multidisciplinary approach combining paleontological, archeological, historical, fisheries, and ecological data to reconstruct past changes in marine populations, habitats, and water quality in the Adriatic Sea. Then, we constructed binary food webs for different historical periods to analyze possible changes in food-web structure and functioning over time. Our results indicate that human activities have influenced marine resource abundance since at least Roman times and accelerated in the nineteenth and twentieth centuries. Today, 98% of traditional marine resources are depleted to less than 50% of former abundance, with large (>1 m) predators and consumers being most affected. With 37% of investigated species rare and 11% extirpated, diversity has shifted towards smaller, lower trophic-level species, further aggravated by more than 40 species invasions. Species providing habitat and filter functions have been reduced by 75%, contributing to the degradation of water quality and increased eutrophication. Increased exploitation and functional extinctions have altered and simplified food-web structure over time, especially by changing the proportions of top predators, intermediate consumers, and basal species. Moreover, simulations of species losses indicate that today’s ecosystems may be less robust to species extinctions than in the past. Our results illustrate the long-term and far-reaching consequences human activities can have on marine food webs and ecosystems.     

Abundance and Distribution of Dimethylsulfoniopropionate Degradation Genes and the Corresponding Bacterial Community Structure at Dimethyl Sulfide Hot Spots in the Tropical and Subtropical Pacific Ocean

Dimethylsulfoniopropionate (DMSP) is mainly produced by marine phytoplankton but is released into the microbial food web and degraded by marine bacteria to dimethyl sulfide (DMS) and other products. To reveal the abundance and distribution of bacterial DMSP degradation genes and the corresponding bacterial communities in relation to DMS and DMSP concentrations in seawater, we collected surface seawater samples from DMS hot spot sites during a cruise across the Pacific Ocean. We analyzed the genes encoding DMSP lyase (dddP) and DMSP demethylase (dmdA), which are responsible for the transformation of DMSP to DMS and DMSP assimilation, respectively. The averaged abundance (±standard deviation) of these DMSP degradation genes relative to that of the 16S rRNA genes was 33% ± 12%. The abundances of these genes showed large spatial variations. dddP genes showed more variation in abundances than dmdA genes. Multidimensional analysis based on the abundances of DMSP degradation genes and environmental factors revealed that the distribution pattern of these genes was influenced by chlorophyll a concentrations and temperatures. dddP genes, dmdA subclade C/2 genes, and dmdA subclade D genes exhibited significant correlations with the marine Roseobacter clade, SAR11 subgroup Ib, and SAR11 subgroup Ia, respectively. SAR11 subgroups Ia and Ib, which possessed dmdA genes, were suggested to be the main potential DMSP consumers. The Roseobacter clade members possessing dddP genes in oligotrophic subtropical regions were possible DMS producers. These results suggest that DMSP degradation genes are abundant and widely distributed in the surface seawater and that the marine bacteria possessing these genes influence the degradation of DMSP and regulate the emissions of DMS in subtropical gyres of the Pacific Ocean.     

Phytoplankton across Tropical and Subtropical Regions of the Atlantic,Indian and Pacific Oceans

We examine the large-scale distribution patterns of the nano- and microphytoplankton collected from 145 oceanic stations, at 3 m depth, the 20% light level and the depth of the subsurface chlorophyll maximum, during the Malaspina-2010 Expedition (December 2010-July 2011), which covered 15 biogeographical provinces across the Atlantic, Indian and Pacific oceans, between 35°N and 40°S. In general, the water column was stratified, the surface layers were nutrient-poor and the nano- and microplankton (hereafter phytoplankton, for simplicity, although it included also heterotrophic protists) community was dominated by dinoflagellates, other flagellates and coccolithophores, while the contribution of diatoms was only important in zones with shallow nutriclines such as the equatorial upwelling regions. We applied a principal component analysis to the correlation matrix among the abundances (after logarithmic transform) of the 76 most frequent taxa to synthesize the information contained in the phytoplankton data set. The main trends of variability identified consisted of: 1) A contrast between the community composition of the upper and the lower parts of the euphotic zone, expressed respectively by positive or negative scores of the first principal component, which was positively correlated with taxa such as the dinoflagellates Oxytoxum minutum and Scrippsiella spp., and the coccolithophores Discosphaera tubifera and Syracosphaera pulchra (HOL and HET), and negatively correlated with taxa like Ophiaster hydroideus (coccolithophore) and several diatoms, 2) a general abundance gradient between phytoplankton-rich regions with high abundances of dinoflagellate, coccolithophore and ciliate taxa, and phytoplankton-poor regions (second principal component), 3) differences in dominant phytoplankton and ciliate taxa among the Atlantic, the Indian and the Pacific oceans (third principal component) and 4) the occurrence of a diatom-dominated assemblage (the fourth principal component assemblage), including several pennate taxa, Planktoniella sol, Hemiaulus hauckii and Pseudo-nitzschia spp., in the divergence regions. Our findings indicate that consistent assemblages of co-occurring phytoplankton taxa can be identified and that their distribution is best explained by a combination in different degrees of both environmental and historical influences.