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: 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 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: 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: In this study, the impact of oceanic data assimilation on ENSO simulations and predictions is investigated. The authors’ main objective is to compare the impact of the assimilation of sea level observations and three-dimensional temperature measurements relative to each other. Three experiments were performed. In a control run the ocean model was forced with observed winds only, and in two assimilation runs three-dimensional temperatures and sea levels were assimilated one by one. The root-mean-square differences between the model solution and observations were computed and heat content anomalies of the upper 275 m compared to each other. Three ensembles of ENSO forecasts were performed additionally to investigate the impact of data assimilation on ENSO predictions. In a control ensemble a hybrid coupled ocean–atmosphere model was initialized with observed winds only, while either three-dimensional temperatures or sea level data were assimilated during the initialization phase in two additional forecast ensembles. The predicted sea surface temperature anomalies were averaged over the eastern equatorial Pacific and compared to observations. Two different objective skill measures were computed to evaluate the impact of data assimilation on ENSO forecasts. The authors’ experiments indicate that sea level observations contain useful information and that this information can be inserted successfully into an oceanic general circulation model. It is inferred from the forecast ensembles that the benefit of sea level and temperature assimilation is comparable. However, the positive impact of sea level assimilation could be shown more clearly when the forecasted temperature differences rather than the temperature anomalies themselves were compared with observations.

Description: The seasonal cycle over the tropical Pacific simulated by 11 coupled ocean–atmosphere general circulation models (GCMs) is examined. Each model consists of a high-resolution ocean GCM of either the tropical Pacific or near-global means coupled to a moderate- or high-resolution atmospheric GCM, without the use of flux correction. The seasonal behavior of sea surface temperature (SST) and eastern Pacific rainfall is presented for each model. The results show that current state-of-the-art coupled GCMs share important successes and troublesome systematic errors. All 11 models are able to simulate the mean zonal gradient in SST at the equator over the central Pacific. The simulated equatorial cold tongue generally tends to be too strong, too narrow, and extend too far west. SSTs are generally too warm in a broad region west of Peru and in a band near 10°S. This is accompanied in some models by a double intertropical convergence zone (ITCZ) straddling the equator over the eastern Pacific, and in others by an ITCZ that migrates across the equator with the seasons; neither behavior is realistic. There is considerable spread in the simulated seasonal cycles of equatorial SST in the eastern Pacific. Some simulations do capture the annual harmonic quite realistically, although the seasonal cold tongue tends to appear prematurely. Others overestimate the amplitude of the semiannual harmonic. Nonetheless, the results constitute a marked improvement over the simulations of only a few years ago when serious climate drift was still widespread and simulated zonal gradients of SST along the equator were often very weak.

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: In this paper the performance of the global coupled general circulation model (CGCM) ECHO-2, which was integrated for 10 years without the application of flux correction, is described. Although the integration is rather short, strong and weak points of this CGCM can be clearly identified, especially in view of the model's performance of the annual cycle in the tropical Pacific. The latter is simulated with more success relative to the earlier version, ECHO-I. A better representation of the low-level stratus clouds in the atmosphere model associated with a reduction in the shortwave radiative flux at the air-sea interface improved the coupled model's performance in the southeastern tropical oceans, with a strongly reduced warm bias in these regions. Modifications in the atmospheric convection scheme also eliminated the AGCM's tendency to simulate a double ITCZ, and this behavior is maintained in the CGCM simulation. Finally, a new numerical scheme for active tracer advection in the ocean model strongly reduced the numerical mixing, which seems to enhance considerably the level of interannual variability in the equatorial Pacific. One weak point is an overall cold bias in the Tropics and midlatitudes, which typically amounts to 1°C in open ocean regions. Another weak point is the still too strong equatorial cold tongue, which penetrates too far into the western equatorial Pacific. Although this model deficiency is not as pronounced as in ECHO-1, the too strong cold tongue reduces the level of interannual rainfall variability in the western and central equatorial Pacific. Finally, the interannual fluctuations in equatorial Pacific sea surface temperatures (SSTs) are too equatorially trapped, a problem that is also found in ocean-only simulations. Overall, however, the authors believe that the ECHO-2 CGCM has been considerably improved relative to ECHO-1.