ALBERT

Description: In both terrestrial and aquatic environments introductions of non-indigenous species are continuing and represent one important component of global change. Negative biotic interactions by resident species may prevent successful invaders from becoming pests. Few experimental data are available on the presence and significance of such biotic resistance other than predation or competition. This study addresses the role of habitat structure provided by a native eelgrass (Zostera marina) canopy on growth and survival of the non-indigenous mussel Musculista senhousia, a habitat-modifying gregarious suspension feeder with strong effects on native infauna and eelgrass. In 2 southern California bays, a series of transplantation experiments using tagged mussels revealed that inside an eelgrass canopy, Musculista growth rates were reduced by more than half in 3 of 4 experiments compared to adjacent unvegetated areas. Musculista survival also decreased inside the vegetation in a 4-mo experiment. As one element of habitat structure, we tested the effects of eelgrass patch size, using natural (1 site) and planted (1 site) eelgrass patches of defined sizes. Growth rates of Musculista were highest outside the vegetation and decreased as eelgrass patch size increased. As a potential mechanism for the canopy effects, we suggest that Musculista receives less food inside the vegetation. In the experimental plots, the presence and spatial extent of the macrophyte canopy strongly affected near bottom (10 cm) horizontal water flow assessed with a direct dye tracking method. Reduced mussel growth rates were linearly associated with lower water flow, and presumably, food flux. Over a period of 7 mo, food resources (particulate chlorophyll a) were consistently lower 1 and 5 cm above the sea floor inside eelgrass patches compared to the sand flat. The reduction in food availability matched the growth reduction of Musculista. Also, mussel condition (dry flesh mass/shell mass) was worse in individuals growing in eelgrass than in the sand flat. Previous experiments revealed that dense beds of Musculista impede the rhizome growth and vegetative propagation of eelgrass, yet mussels attain abundances sufficient for interference only if eelgrass beds are patchy. Thus, anthropogenic disturbances on eelgrass beds, which often result in meadow fragmentation, and the proliferation of Musculista may have synergistic negative effects on the persistence of eelgrass beds.

Description: In situ experiments were run with the seastar Asterias rubens to investigate the influence of epibiosis on predation preferences. Mussels (Mytilus edulis) monospecifically fouled by different epibiont species (the barnacle Balanus improvisus, the red filamentous alga Ceramium strictum, the sponge Halichondria panicea and the hydrozoan Laomedea flexuosa) and macroscopically clean mussels were exposed and seastar predation was monitored by SCUBA. Asterias rubens preferred macroscopical unfouled mussels as prey. Fouling generally reduced predation pressure on the mussel hosts (associational resistance). Barnacles protected mussels less efficiently than hydrozoans or algae. We hypothesize that in top-down controlled communities this influence of epibiosis on predation pressure should affect mussel community patterns. A survey of natural mussel-epibiont distribution in the presence or absence of A. rubens showed that the prevalence of differently fouled mussels differed between predation-exposed and predation-protected habitats. Natural mussel-epibiont associations reflected the preferential predation of the major local predators. Additionally, higher epibiotic diversity and evenness could be observed at locations accessible to benthic predators as compared with habitats protected from predation. As blue mussels and seastars are important structuring and controlling elements in the shallow water community of Kiel Fjord, major consequences of epibiosis on the entire system are discussed.

Description: The El Niño/Southern Oscillation (ENSO) phenomenon is the strongest natural interannual climate fluctuation1. ENSO originates in the tropical Pacific Ocean and has large effects on the ecology of the region, but it also influences the entire global climate system and affects the societies and economies of manycountries2. ENSO can be understood as an irregular low-frequency oscillation between a warm (El Niño) and a cold (La Niña) state. The strong El Niños of 1982/1983 and 1997/1998, along with the more frequent occurrences of El Niños during the past few decades, raise the question of whether human-induced 'greenhouse' warming affects, or will affect, ENSO3. Several global climate models have been applied to transient greenhouse-gas-induced warming simulations to address this question4, 6, but the results have been debated owing to the inability of the models to fully simulate ENSO (because of their coarse equatorial resolution)7. Here we present results from a global climate model with sufficient resolution in the tropics to adequately represent the narrow equatorial upwelling and low-frequency waves. When the model is forced by a realistic future scenario of increasing greenhouse-gas concentrations, more frequent El-Niño-like conditions and stronger cold events in the tropical Pacific Ocean result

Description: The overflow and descent of cold dense water from the Denmark Strait sill-a submarine passage between Greenland and Iceland-is a principal means by which the deep ocean is ventilated, and is an important element in the global thermohaline circulation. Previous investigations of its variability-in particular, direct current measurements(1,2) in the overflow core since 1986-have shown surprisingly little evidence of long-term changes in now speed. Here we report significant changes in the overflow characteristics during the winter of 1996-97, measured using two current-meter moorings and an inverted echo sounder located at different depths in the fastest part of the now. The overflow warmed to the highest monthly value yet recorded (2.4 degrees C), and showed a pronounced slowing and thinning at its lower margin. We believe that the extreme warmth of the overflow caused it to run higher on the continental slope off east Greenland, so that the lower current meters and the echo sounder were temporarily outside and deeper than the fast-flowing core; model simulations appear to confirm this interpretation, We suggest that the extreme warmth of the overflow is a lagged response to a warming upstream in the Fram Strait three years earlier (caused by an exceptional amplification of the winter North Atlantic Oscillation). If this is so, over-now characteristics may be predictable.

Description: The use of stable isotope analysis in ecological and wildlife studies is rapidly increasing. Studies include evaluating flow of nutrients in ecosystems and studying dietary composition of individual animals. Several mixing models have been developed to evaluate the relative contribution of different foods to the diet of consumers. All these mixing models require that all prey types will be significantly different in bivariate space. This requirement usually poses a problem in analyzing data of stable isotope ratios because sample sizes in most studies are small and seldom normally distributed. We propose a randomization test that we based on the K nearest-neighbor approach. Results from our simulations of power revealed that the K nearest-neighbor test appears to have high power even with small sample sizes and comparatively low displacement. The K nearest-neighbor test described here provides the preliminary statistical analysis necessary for the use of the mixing models, and therefore is a new, powerful tool for analyzing stable isotope data. In evaluating the test performance on data collected from American martens (Martes americana) and their prey on Chichagof Island, Southeast Alaska, we were able to reject our null hypothesis that all samples of prey were drawn from identical populations (P = 0.05). A program written in Pascal or S-Plus is available from the authors to evaluate the K nearest-neighbor statistic for several groups.

Description: In steady state, the export of photosynthetically fixed organic matter to the deep ocean has to be balanced by an upward flux of nutrients into the euphotic zone1. Indirect geochemical estimates2 of the nutrient supply to surface waters have been substantially higher than direct biological and physical measurements3, particularly in subtropical regions. A possible explanation for the apparent discrepancy is that the sampling strategy of the direct measurements has under-represented episodic nutrient injections forced by mesoscale eddy dynamics, whereas geochemical tracer budgets integrate fluxes over longer time and space scales. Here we investigate the eddy-induced nutrient supply by combining two methods potentially capable of delivering synoptic descriptions of the ocean's state on a basin scale. Remotely sensed sea-surface height data from the simultaneous TOPEX/Poseidon and ERS-1 satellite missions are assimilated into a numerical eddy-resolving coupled ecosystem–circulation model of the North Atlantic Ocean. Our results indicate that mesoscale eddy activity accounts for about one-third of the total flux of nitrate into the euphotic zone (taken to represent new production) in the subtropics and at mid-latitudes. This contribution is not sufficient to maintain the observed primary production in parts of the subtropical gyre, where alternative routes of nitrogen supply will have to be considered.

Description: Two sets of equations, covering all world oceans and seas, are presented to calculate pressure from depth for the computation of sound speed, and depth from pressure for use in ocean engineering. They are based on the algorithm of UNESCO 1983 [N. P. Fofonoff and R. C. Millard, Jr., Unesco Tech. Papers in Mar. Sci. No. 44 (1983)], and on calculations from temperature and salinity profiles. The pressure to depth conversion is presented first. The equations can be used in those cases where the desired accuracy is reduced to ±0.8 m. The equations to convert depth to pressure provide an overall accuracy between ±8000 Pa and ±1000 Pa. This leads to errors in sound speed consistently smaller than ±0.02 m/s. The discussion, and comparisons with results and other formulas, suggest that the new equations are a substantial improvement on the previous simplified ones, which should now be abandoned.

Description: The ability to monitor the heat content of oceans over long distances is becoming increasingly important for understanding the role of oceans in climate change, for determining the variability of the state of the oceans, for operational ocean observing systems, and for studying large-scale ocean processes such as water-mass formation. Although the properties of the upper layers of the ocean can be routinely measured on large scales by satellite remote sensing (providing altimetric and infrared data) and with expendable probes dropped from commercial vessels, the deep interior of the ocean is more difficult to monitor. Ocean acoustic tomography1 is a promising technique for such applications, as it has the potential to provide systematic, instantaneous and repeated measurements of the ocean interior over large parts of an ocean basin. Here we demonstrate the capability of this technique for measuring the heat content across an entire (albeit small) ocean basin—the western Mediterranean Sea.

Description: Newly hatched Baltic cod Gadus morhua larvae are typically found at depths 〉60 m. This is a region of low light and prey availability, hence generating the hypothesis that larvae have to migrate from hatching depth to the surface layer to avoid starvation and improve their nutritional condition. To test this hypothesis, Baltic cod larvae were sampled during the spawning seasons of 1994 and 1995 with depth-resolving multiple opening/closing nets. Each larva was aged by otolith readings and its RNA/DNA ratio was determined as a measure of nutritional condition. The RNA/DNA ratios of these larvae aged 2-25 days (median 10 days) ranged from 0.4 to 6.2, corresponding to levels exhibited by starving and fast-growing larvae in laboratory calibration studies (starvation, protein growth rate, Gpi= -12.2% day−1; fastgrowing larvae, Gpi=14.1%day−1) respectively. Seventy per cent of the field caught larvae had RNA/DNA ratios between the mean values found for starving and fed laboratory larvae. Only larvae aged 8-11 days had higher mean RNA/DNA ratios above 45 m than below (t-test, P〈0.05). However, the instantaneous protein growth rates were significantly higher for all larval age groups in the surface layers (t-test, P〈0.05). Starving larvae were found in all depths sampled (10-85 m), whereas growing larvae (positive Gpi) were restricted to samples taken shallower than 45 m. These superior growth rates above 45 m corroborate the hypothesis and imply that migration to the shallow water layers is a prerequisite for good nutritional condition, growth and survival of Baltic cod larvae. The frequent occurrence of cod larvae older than 8 days in the deep water in poor condition suggests that a proportion of the larvae will die from Starvation in the deep layers of the Baltic Sea.

Description: The ocean stores and transports vast quantities of heat, fresh water, carbon and other materials, and its circulation plays an important role in determining both the Earth's climate and fundamental processes in the biosphere. Understanding the development of climate and important biological cycles therefore requires detailed knowledge of ocean circulation and its transport properties. This cannot be achieved solely through modelling, but must involve accurate observations of the spatio-temporal evolution of the global oceanic flow field. Estimates of oceanic flow are currently made on the basis of space-borne measurements of the sea surface, and monitoring of the ocean interior. Satellite altimetry and acoustic tomography are complementary for this purpose1, as the former provides detailed horizontal coverage of the surface, and the latter the requisite vertical sampling of the interior. High-quality acoustic-tomographic2 and altimetric3 data are now available to test the combined power of these technologies for estimating oceanic flows. Here we demonstrate that, with the aid of state-of-the-art numerical models, it is possible to recover from these data a detailed spatio-temporal record of flow over basin-scale volumes of fluid. Our present results are restricted to the Mediterranean Sea, but the method described here provides a powerful tool for studying oceanic circulation worldwide.

Description: Recent captures of two female giant squid ( Architeuthis ) off southern Australia have provided the first record of a mated female specimen of these almost mythical deepsea creatures. We found sperm packages (spermatophores) embedded within the skin of both ventral arms of the larger of the two specimens. It seems that male giant squids may use their muscular elongate penis to ‘inject’ sperm packages under pressure directly into the arms of females.

Description: The nitrogen-isotope record preserved in Southern Ocean sediments, along with several geochemical tracers for the settling fluxes of biogenic matter, reveals patterns of past nutrient supply to phytoplankton and surface-water stratification in this oceanic region. Areal averaging of these spatial patterns indicates that reduction of the CO2 'leak' from ocean to atmosphere by increased surface-water stratification south of the Polar Front made a greater contribution to the lowering of atmospheric CO2 concentration during the Last Glacial Maximum than did the increased export of organic carbon from surface to deep waters occurring further north.

Description: The conventional model whereby plume volcanism forms linear age-progressive volcanic chains, with the youngest activity occurring nearest a spreading axis (at a 'hotspot'), has been challenged for the Easter seamount chain1–4. Whereas early work suggested the existence of a linear melting anomaly (a 'hotline')1,2, more recent studies3,4 have proposed a hotspot near Salas y Gomez island, connected with the Easter microplate spreading system by an ~800-km-long, volcanically active plume channel. Here we use geochemical, geological and geochronological data to argue that the hotspot lies close to Easter Island. Moreover, new isotopic data for lavas from the seamount chain provide evidence for bidirectional flow between the spreading axis and the plume, thus supporting geophysical and fluid-dynamical models of mantle flow in a plume/spreading axis system5–7. Material balance and flux considerations show the Easter plume to be weak and cool compared with those beneath larger features such as Iceland, Hawaii and the Galápagos islands.

Description: A knowledge of past changes in the biological productivity of the oceans is important for understanding the interactions between carbon cycling and climate. Phytoplankton productivity in today's oceans can be estimated from the concentrations of chlorophyll in sea water1, but chlorophyll is not preserved in the sediments. Existing proxies for past algal productivity do not represent total productivity; for example, biogenic opal2 reflects the contribution of only part of the phytoplankton community, and the organic carbon record can be subject to contamination from terrestrial inputs2,3. Although chlorins, the pigment-transformation products of chlorophyll, are widespread in Quaternary marine sediments, their potential as proxy measures of past variations in primary productivity has not been convincingly demonstrated. Here we report a high-resolution molecular stratigraphic record of chlorin concentrations over the past 350,000 years in a sediment core from the subtropical Atlantic continental margin. Maxima in the chlorin accumulation rate coincide with significant peaks in the accumulation rates of biogenic opal (at the end of glacial terminations) and organic carbon (between terminations). These results suggest that chlorins, unlike other proxies, can serve as a measure of total primary productivity variations.

Description: Attributes have proliferated recently with different selections available on different workstations. What do they all mean? When do we use one and when another? The answers to these questions are not easy but the first step is to understand what our options are, and herein lies the purpose of this article.

Description: A fundamental issue in marine science is the identification of the factors controlling biological uptake of CO2, in high-nitrate, low-chlorophyll regions. A recent in situ iron fertilization experiment demonstrated that iron limitation is responsible for low phytoplankton stocks in the equatorial Pacific4. Here we show that flavodoxin, a biochemical marker of iron limitation, can be used to map the degree of iron stress in natural populations. Flavodoxin assays along a 900-km east-west transect in the northeastern subarctic Pacific revealed a pronounced increase in iron stress in the region west of the 135° W meridian. Addition of dissolved iron alleviated this stress. Immunostaining of single cells from the most western station showed that flavodoxin is present specifically within the chloroplasts of diatoms. Our approach provides a rapid means of defining the extent of iron stress in the ocean5 and supports the hypothesis that diatoms are iron stressed in the northeast Pacific.

Description: We propose that ocean conditions of the Near Islands in the western Atleutian Arc mimic those of the shallow continental shelf of the eastern Bering Sea to the extent that the marine community, including assemblages of forage fishes and their avian predators, has disinctly coastal characteristics. In contrast, marine avifauna and their prey at neighbouring Buldir Island are distinctly oceanic. For example, at the Near Islands, the ratio of thick-billed to common murres, Uria lomvia and U. aalge, is low and black.legged kittiwakes, Rissa tridacytla, but not red-legged kittiwakes, R. brevirostris, nest there. Diets of murres and kittiwkaes are dominated by sand lance, Ammodytes hexapterus, an abundant coastal species. At Buldir Island, thick-billed murres greatly outnumber common murres, red-legged kittiwakes and black-legged kittiwakes are both abundant, and diets of the birds consist primarily of oceanic squid and lantern-fish (Myctophidae). This mesoscale difference in food webs is apparently a consequence of the local physiography. A broad escarpment on the Near physiographic block creates a comparatively expansive, shallow, shelf-like habitat around the Near Islands, where a pelagic community typical of coastal regions flourished. Buldir Island is the only emergent feature of the Buldir physiographic block, with little shallow water surrounding it and, apparently, little opportunity for other than oceanic species to exist. Patterns in the distribution of fishes, and thus of sea birds, throughout the Atleutian Islands might be largely explained by the relationship between physical environments and food webs. In the larger context of fisheries oceanography, this model for the Aleutian Islands improves our ability to interpret physical and biological heterogeneity in the ocean and its relationship to regional community dynamics and trends in the abundance and productivity of individual species at higher tropic levels.

Description: The temperature relationship of routine metabolic rate (Rr) of non-feeding, non-growing Coregonus lavaretus larvae between 2 and 15°C is characterized by Q10-values ranging from l.8-2.45. The rate of growth, based on weight determinations, of first-feeding larvae amounted to 3.5, 7.6 and 9.4% day-1 at 5, 10 and 12°C respectively, from which Q10-values between 4.0 and 4.8 can be calculated. The rate of increase of muscle mass between 5 and 10°C, based on the determination of the cross-sectional area of inner muscle fibres, resulted in a Q10-value of 4.5. Water temperature influenced the pattern of growth of the inner muscle fibres. At hatching, after 360 day degrees, total muscle mass of larvae reared at 4 and 8°C was independent of temperature, but at 4°C the rate of mass increase owed more to hyperplasia (increase in fibre number) than to hypertrophy (increase in fibre mass), whereas at 8°C the opposite was the case. The calculation of power budgets (including the metabolic cost of growth) of first-feeding larvae yielded net conversion efficiencies (K2) increasing with temperature from 46.3% at 5°C to 54.7% at 12°C. Comparing our data with literature data two general conclusions can be drawn. (1) In first-feeding larvae the net, but not the gross, conversion efficiency of food energy increases with temperature. This is due to net energy input being characterized by a much higher Q10-value than energy expenditures. (2) In embryos of freshwater fish so far investigated hyperplasia plays a greater role in the increase of fibre mass than hypertrophy at the lower temperature, whereas in embryos of marine fish hyperplasia prevails at the higher temperature. It is suggested that this discrepancy correlates with the high concentration of free amino acids in the eggs of marine species which provide an additional, easily available, source of metabolic energy absent in freshwater species.

Description: Seismic data are usually acquired and processed for imaging reflections. This paper describes a method of processing seismic data for imaging discontinuities (e.g., faults and stratigraphic features). One application of this nontraditional process is a 3-D volume, or cube, of coherence coefficients within which faults are revealed as numerically separated surfaces. Figure 1 compares a traditional 3-D reflection amplitude time slice with the results of the new method. To our knowledge, this is the first published method of revealing fault surfaces within a 3-D volume for which no fault reflections have been recorded.

Description: Seismic tomography and the isotope geochemistry of Cenozoic volcanic rocks suggest the existence of a large, sheet-like region of upwelling in the upper mantle which extends from the eastern Atlantic Ocean to central Europe and the western Mediterranean. A belt of extension and rifting in the latter two areas appears to lie above the intersection of the centre of the upwelling region with the base of the lithosphere. Lead, strontium and neodymium isotope data for all three regions converge on a restricted composition, inferred to be that of the upwelling mantle.

Description: THE hydrothermal circulation of sea water through permeable ocean crust results in rock–water interactions that lead to the formation of massive sulphide deposits. These are the modern analogues of many ancient ophiolite-hosted deposits1–4, such as those exposed in Cyprus. Here we report results obtained from drilling a series of holes into an actively forming sulphide deposit on the Mid-Atlantic Ridge. A complex assemblage of sulphide–anhydrite–silica breccias provides striking evidence that such hydrothermal mounds do not grow simply by the accumulation of sulphides on the sea floor. Indeed, the deposit grows largely as an in situ breccia pile, as successive episodes of hydrothermal activity each form new hydrothermal precipitates and cement earlier deposits. During inactive periods, the collapse of sulphide chimneys, dissolution of anhydrite, and disruption by faulting cause brecciation of the deposit. The abundance of anhydrite beneath the present region of focused hydrothermal venting reflects the high temperatures ( 〉 150 °C) currently maintained within the mound, and implies substantial entrainment of cold sea water into the interior of the deposit. These observations demonstrate the important role of anhydrite in the growth of massive sulphide deposits, despite its absence in those preserved on land.