ALBERT

Beschreibung: Latitudinal declines of species richness from the tropics to the poles represent a general spatial pattern of diversity on land. For the marine realm, the generality of this pattern has frequently been questioned. Here, I use a database with nearly 600 published gradients (198 of which were marine) to assess whether there is a marine latitudinal diversity gradient of similar average strength and slope as that for terrestrial organisms. Using meta-analysis techniques, I also tested which characteristics of organisms or habitats affected gradient strength and slope. The overall strength and slope of the gradient for marine organisms was significantly negative and of similar magnitude compared to gradients for terrestrial organisms. Marine gradients were on average stronger as well as steeper than freshwater gradients. Latitudinal gradients were clearly a regional phenomenon, with stronger gradients and steeper slopes for diversity assessed on regional than on local scales. The gradient parameters differed also between oceans and between different habitats, with steeper gradients related to the pelagial rather than the benthos. There were on the other hand no significant differences between hemispheres and between different gradient ranges, although such differences have often been presumed. The most important organismal characteristic related to gradient structure was body mass, with significant gradients related to large organisms. A significant increase in gradient strength with increasing trophic level was observed. The meta-analysis also revealed strongest gradients for nekton and mobile epifauna, whereas the gradients were weak for sessile epifauna and for infauna. In conclusion, marine biota reveal a similar overall decline in diversity with latitude to that observed in terrestrial realms, but the strength and slope of the gradient are clearly subject to regional, habitat and organismal features.

Beschreibung: Grazing experiments were conducted with natural mesozooplankton from Kiel Bight, Germany, using radioactive labelled phytoplankton cultures and seston size fractions. The results of experiments using phytoplankton cultures indicated that bivalve veligers performed highest clearance of particles within a size range of 4.7 to 6.3 µm, whereas optimum particle size for copepods was 15 µm. The results of experiments using labelled natural seston size fractions identified bivalve veligers and appendicularians as those responsible for the removal of particles within the smallest size class (〈2 µm). Seston size fractions larger than 5 µm were mainly cleared by copepods and nauplii. As particle size increased, the contribution of copepod clearance to total zooplankton clearance within size classes increased from 57% (〈5 µm size class) to more than 81% (30 to 100 µm size class). When the nauplii clearance rates were included, the total copepod clearance accounted for 90 to 97.6% of the total volume cleared of particles bigger than 10 µm. Despite low abundances of bivalve veligers and appendicularians in Kiel Bight at the time of the experiment, we calculated that approximately 10 and 8.5%, respectively, of the carbon ingested by total mesozooplankton was due to veliger and appendicularian grazing. The importance of bivalve veligers might be seen in their grazing on seston particles that escape predation by copepods and on the amount of energy that is therefore directed from the water column to the benthos when larvae settle.

Beschreibung: Filamentous sulfur-oxidizing bacteria and geochemical parameters of sediments at the Makran accretionary wedge in the northeastern Arabian Sea off Pakistan were studied. The upper continental slope between 350 and 850 m water depth, which is in the center of the oxygen-minimum zone, is characterized by numerous sites of small-scale seeps of methane- and sulfide-charged porewater. White bacterial mats with diameters 〈1 m were discovered at the surface of these sites using a photo-TV sled. Seep sediments, as well as non-seep sediments, in the vicinity were characterized by the occurrence of the bacterium Thioploca in near-surface layers between 0 and 13 cm depth. Thioploca bundles were up to 20 mm in length and contained up to 20 filaments of varying diameters, between 3 and 75 µm. Up to 169 ind. cm-2 were counted. Maximum numbers occurred in the top 9 cm of sediment, which contained very low concentrations of soluble sulfide (〈0.2 µM) and high amounts of elemental sulfur (up to 10 µmol cm-3). Moderate sulfate reduction activity (between 20 and 190 nmol cm-3 d-1) was detected in the top 10 cm of these sediments, resulting in a gradual downcore decrease of sulfate concentrations. CO2 fixation rates had distinct maxima at the sediment surface and declined to background values below 5 cm depth. The nutritional implications of the distinct morphology of Thioploca and of the geochemical setting are discussed and compared to other sites containing Thioploca communities.

Beschreibung: Four in situ experiments were conducted to examine the potential top-down and bottom-up control of epibenthic ciliate communities. The experiments were run in the littoral of Lake Erken and at a brackish water site on the island of Väddö on the Baltic coast of Sweden, during the spring of 2000. The experimental manipulations were the presence/absence of the natural macrozoobenthos grazer community, cross-classified with the presence/absence of additional nutrients. Epibenthic ciliates responded to both manipulation of grazers and resources, but the response was group specific. Total ciliate abundance decreased when macrozoobenthos (largely chironomids, gastropods, trichopteran larvae, isopods and amphipods) were removed, thus excluding a direct predation effect of the macrozoobenthos community on ciliates. Total ciliate biomass, but not abundance, tended to increase in the presence of additional nutrients; an effect weakly dependent on season and site. The disparity between effects of nutrients on biomass and abundance was due to effects on heterotrichs, a group of large but relatively rare algivorous ciliates. The manipulations altered the ciliate community composition, and between lakes there were differences in species richness and diversity and experiments. However, neither the removal of macrozoobenthos nor the addition of nutrients changed species richness or diversity. This runs counter to work with other taxonomic groups, which shows maximal diversity at an intermediate level of resources or predation. This can only be partially explained by the lack of direct predation effects and the open nature of the experimental system.

Beschreibung: Spatial and temporal variability in environmental factors can exert major influences on survival and growth of living organisms. However, in many key areas of fisheries science (e.g. growth, survival and recruitment determination), environmental heterogeneity is usually ignored because of insufficient environmental or fisheries data or lack of evidence that such heterogeneity impacts response variables. For the eastern Baltic Sea (ICES Subdivisions 25 to 32), we evaluated spatial and temporal differences in conditions affecting the survival of cod Gadus morhua L. eggs at survival on four distinct spawning sites within the assessment area. We intercalibrated ways of quantifying the volume of water ('reproductive volume') at each site where salinity, oxygen and temperature conditions permitted successful egg development. We have developed and compared a time series (1952 to 1996) of reproductive volumes among the areas to identify spatial differences. The results of 2 independent volume-estimation methods are comparable, indicating that highly significant differences exist among the sites, and that the westernmost spawning ground, Bornholm Basin, has on average the highest reproductive volume and the lowest variability among the 4 sites. These findings may be useful in evaluating how spatial and temporal variability in environmental conditions affect egg hatching success and possibly recruitment in the Baltic stock.

Beschreibung: Natural marine bacteria populations collected from nearshore waters produce different types of siderophores depending on the degree of iron limitation. These siderophores can facilitate iron uptake in the marine diatom Phaeodactylum tricornutum. Water samples from 15 stations along the Italian coast of the northwest Adriatic Sea were collected and filter fractionated (3.0, 0.8 and 0.2 µm). Siderophore production in the fractions was determined using cross-feeding experiments with siderophore-auxotrophic bacteria. At most stations sampled, bacteria collected in the 3.0 and 0.8 µm filters produced siderophores which stimulated growth in Morganella morganii, the indicator strain for α-keto/ α-hydroxy acids. The results suggest that MGF (ŒMorganella-Growth Factor¹) production is common among filamentous and appendaged bacteria or strains associated with particles. Natural bacteria populations grown in iron-deficient media stimulated growth of all the indicator strains in the cross-feeding tests. Examples of known MGF which supply iron to M. morganii were tested for their ability to act as iron source for the marine diatom P. tricornutum. Iron uptake from 55Fe-MGFs was measured in P. tricornutum cells grown in Fe-sufficient and Fe-deficient media. Unchelated iron (55FeCl3 ) and 55FeEDTA were used as controls. The uptake of iron from the 55Fe-MGF and 55FeCl3 by Fe-deficient cells was higher (109 to 150 pgFe mg-1) than from 55FeEDTA (34 pgFe mg-1). Similarly, Fe-sufficient P. tricornutum took up iron from the 55Fe-MGF and 55FeCl3 to the same extent (~50 pgFe mg-1) while minimal uptake (8 pgFe mg-1) was measured from FeEDTA. In growth experiments where iron-deficient diatom cells were incubated in media containing different sources of iron, e.g. FeCl3, Fe-MGF and FeEDTA, a greater increase in number was observed in cells supplied with Fe-MGF. Further experiments also show that the uptake of Fe from MGF was enhanced by light and that a reduction step was involved in the uptake process. MGF also promoted the uptake of colloidal ferrihydrites. This study gives further evidence that siderophores produced by bacteria can be utilized by phytoplankton as an iron source. We therefore suggest that these substances play an important role in increasing the availability of iron to phytoplankton in coastal waters and thus are major factors defining the chemistry of iron in the marine environment.

Beschreibung: Marine angiosperms, or seagrasses, continue to be a major focus of marine biologists because of their important ecological role in many coastal ecosystems. Seagrass population biology could benefit from a population genetic perspective because genetic data enable the extraction of useful demographic information such as isolation and gene flow between demes. Moreover, population genetic processes may contribute to the growing ecological risks of local population extinction. Progress in seagrass genetics is partly driven by novel genetic markers which detect variation at the DNA level and overcome the limited polymorphism of allozymes. Key results of studies in the past decade, mostly using RAPD and microsatellites, were (1) considerable genetic and genotypic (clonal) diversity is present in several species in contrast to earlier notions of low polymorphism detected at allozyme loci, and (2) genetic differentiation among populations seems to be the rule despite earlier reports of genetic uniformity. Pronounced genetic structure was detected between populations of 4 species examined thus far (Posidonia oceanica, P. australis, Zostera marina, Thalassia testudinum). The FST estimates varied widely and ranged from 0.01 to 0.623 across studies and species. Genetic differentiation at a systematic range of scales was only studied in eelgrass Zostera marina, where it was positively correlated with geographic distance. The high polymorphism of RAPD or microsatellite markers will allow the augmention of indirect estimates of gene flow by methods detecting individual immigration events through paternity analysis or assignment tests. Important conservation related issues such as the level of inbreeding and the effective population size have also been obtained from genetic marker data, but results are too scarce at the moment to allow generalizations. In Zostera marina and Posidonia australis, several population genetic attributes such as clonal diversity, mating system and effective population size varied among populations within species, highlighting that there is no Œtypical¹ population. An important gap in our knowledge is whether the effects of natural population fragmentation and patchiness enhance the genetic isolation of populations due to anthropogenic disturbances. It is also unclear whether genetic differentiation displayed at marker loci are correlated with fitness-related plant traits, and whether genetic or genotypic diversity is important for medium- to long-term meadow persistence. An assessment of the genetic and genotypic diversity at marker loci should be combined with experiments on the ecological plasticity and reaction norms of genotypes composing the populations in question. This way, the role of genetic diversity for seagrass population maintenance and growth in the face of changing environmental conditions can be evaluated.

Beschreibung: The epibacterial chemical defense of the marine sponge Suberites domuncula was explored by screening sponge extract, sponge primmorph (3-D aggregates containing proliferating cells) extract and sponge-associated as well as primmorph-associated bacteria for antibacterial activity. 16S rDNA sequencing revealed that the antimicrobially active bacteria belonged to the a- and γ-subdivisions of Proteobacteria ( α-Proteobacterium MBIC 3368, Idiomarina sp. and Pseudomonas sp., respectively). Moreover, a recombinant perforin-like protein was cloned from S. domuncula that displayed strong antibacterial activity. Based on these observations, it is proposed that the sponge may be provided with a direct (by producing antibacterial metabolites) as well as an indirect (with the help of associated bacteria) epibacterial defense.

Beschreibung: There is mounting molecular evidence that bacteria belonging to the phylum Planctomycetes are abundant in marine sponges including members of the genus Aplysina. In an attempt to culture planctomycete bacteria from Aplysina sponges, 116 bacterial strains were isolated on selective oligotrophic media. Screening of the strain collection by fluorescence in situ hybridization with the planctomycete-specific probe Pla46 yielded 3 positive candidates. Nearly complete sequencing of the respective 16S rRNA genes revealed that the isolates were affiliated with 2 distinct clusters of the genus Pirellula: 1 isolate was obtained from a Mediterranean sponge, 1 from a Caribbean sponge and a third from Caribbean seawater. To our knowledge this is the first report of cultured Planctomycetes from marine sponges. The isolates grew slowly on oligotrophic media and failed to grow on nutrient-rich media. Pirellula sp. Strain 797 was pink-pigmented while the other 2 isolates, 16 and 81, were non-pigmented. Transmission electron microscopy revealed a pear- or droplet-shaped cell morphology that is characteristic of the genus Pirellula. The application of strain-specific oligonucleotide probes to sponge tissue cryosections showed that the isolates contribute only a minor fraction to the total microbial community that is associated with Aplysina spp. sponges

Beschreibung: Throughout the last 2 decades a shift from a cod- to a sprat-dominated system occurred in the upper trophic levels of the Central Baltic Sea. This was caused by a decline in the cod stock, due to recruitment failure and high fishing intensity, resulting in a decrease in predation pressure on sprat. Concurrently with the lowest cod stock size on record, sprat reached biomass values of above 2 × 106 t in 1992, being relatively stable afterwards. Besides predation mortality through cod and in recent years also an increasing fishing pressure, cannibalism on eggs may be a compensatory process limiting the reproductive success of sprat and hence contributing to the population regulation in the Central Baltic. Based on sprat stomach sampling on 21 cruises between March 1988 and July 1996 cannibalism on sprat eggs was investigated in the Bornholm Basin, one of the main spawning areas of Central Baltic sprat. Using a model of gastric evacuation to estimate daily food intake rates and a Virtual Population/Extended Survivor Analysis for computing predator population sizes, egg cannibalism rates were estimated. These were compared to egg abundance data from ichthyoplankton surveys and to preliminary estimates of seasonal egg productions. The study revealed significant interannual differences in the intensity of sprat egg cannibalism with considerable predation in 1990 to 1992 (〉15% of the egg abundance during peak spawning and 〉60% of the seasonal production) and a reduction in most recent years (〈16% of the corresponding abundance and production). As a possible reason for these differences a combination of changes in the vertical overlap of predator/prey and variability in the food environment were identified. Shortcomings of the applied methods and the possible impact of cannibalism on the reproductive success and population development of sprat in the Central Baltic Sea are discussed.

Beschreibung: Nanoplankton and picoplankton abundance and community grazing on picoplankton were determined in summer and autumn at several stations in a productive coastal environment (Georges Bank, NW Atlantic Ocean) and in an oligotrophic oceanic ecosystem (Sargasso Sea). Ranges of heterotrophic nanoplankton (HNAN) abundance were 1.2 to 3.6 x 103 cells ml-1 on Georges Bank, and 2.2 to 6.8 x 102 cells ml-1 in the Sargasso Sea. Ranges of phototrophic nanoplankton (PNAN) abundance in these ecosystems were 1.9 to 6.0 x 103 and 1.3 to 4.7 x 102, respectively. Mixotrophic nanoplankton (MNAN), operationally defined here as chloroplast-bearing nanoplankton that ingested fluorescent tracers, comprised an average of 12 to 17% of PNAN in surface waters in both environments during August and October. Mixotrophs at specific stations constituted as much as 38% of total PNAN abundance on Georges Bank and 30% in the Sargasso Sea. Mixotrophs represented up to 39% of the total phagotrophic nanoplankton abundance (MNAN/[MNAN + HNAN]). Community grazing impact was estimated from the disappearance of fluorescent prey surrogates (fluorescently labeled bacteria, FLB; cyanobacteria, FLC; and 〈\3 µm algae, FLA). Absolute grazing rates (total picoplankton cells removed d-1) on Georges Bank exceeded those in the Sargasso Sea due to the greater abundances of predators and prey. However, there was overlap in the specific grazing losses at the 2 sites (ranges = 0.08 to 0.38 d-1 in the coastal ocean and 0.05 to 0.24 d-1 in the oligotrophic ocean). Rates of bacterivory were in approximate balance with rates of bacterial production (3H-thymidine uptake), but production exceeded bacterivory on Georges Bank during the summer cruise. These data are among the first documenting the impact of grazing on picoplankton in these environments, and they are consistent with the prediction that nanoplanktonic protists are major predators of picoplankton. While the proportion of phototrophs that are phagotrophic was highly variable, our study indicates that algal mixotrophy is widespread in the marine environment, occurring in both coastal and oligotrophic sites, and should be considered quantitatively in microbial food web investigations.

Beschreibung: Phytoplankton (〉100 µm) abundance was studied in the open waters of the Gulf of Aqaba during the summer stratification period of 1996. A succession took place among the major phytoplankton groups, with diatom numbers decreasing throughout the summer. The diazotrophic cyanobacteria Trichodesmium spp. became more prominent as the stratification period progressed; 5 Trichodesmium species were identified: T. thiebautii, T. erythraeum with tuft-shaped colonies and Trichodesmium sp. with puff-shaped colonies were common at ~102 colonies m-3 throughout the stratification period, whereas T. tenue and T. hildebrandtii were more rare. A bloom of T. thiebautii and T. erythraeum with 〉106 tuft colonies m-3 was observed in coastal waters of the Gulf during fall 1997. Tuft-shaped colonies were dominant near the surface, while puff-shaped colonies of Trichodesmium sp. were mainly found in the bottom half of the photic zone. These depth distributions were maintained for more than 2 mo, suggesting that the 2 colony types occupied distinct niches. Puff-shaped colonies were found to have higher chlorophyll a contents than tufts, but their photosynthetic activities were not significantly different. Fatty acid analysis of dominant plankton species yielded new trophic relationships for Trichodesmium spp. The Trichodesmium spp.-specific fatty acid C22:2 ω6 was found in Macrosetella gracilis (the sole copepod to graze on Trichodesmium spp.) and in chaetognaths, suggesting that these carnivorous zooplankton fed on M. gracilis. Furthermore, this fatty acid was observed in the filter-feeding Salpa maxima, which was abundantly present in the Gulf of Aqaba during June 1997.

Beschreibung: Nitrogen fixation rates and related plankton parameters were determined at 8 stations in the Baltic proper and Mecklenburg Bay in 1997 and 1998. Nitrogen fixation was measured with 15N-tracer method in unenriched samples. Measurable nitrogen fixation rates were found from July to October, with highest rates in August when heterocystous cyanobacteria formed blooms. Nitrogen fixation rates measured during a moderate bloom in 1998 were related to biomass of heterocystous and coccoid (non-heterocystous) cyanobacteria and primary production. The size fraction 〈10 µm contributed significantly to total nitrogen fixation both during day and night. It is discussed whether this may be due to small, non-heterocystous cyanobacteria, which were abundant in summer and autumn. They may separate photosynthesis and nitrogen fixation temporally (day and night) and may be especially responsible for the high nitrogen fixation rates observed in the dark. As the fraction of pico- and nanoplankton was not considered in earlier studies, a new budget of nitrogen fixation in the Baltic proper has been estimated. On average, daily nitrogen fixation rates of 2.5 mmol N m-2 d-1 (in July/August 1997/1998) and mean annual nitrogen fixation of 125 mmol N m-2 yr-1 were estimated for the Baltic proper. The high variability is discussed. For summer 1998, a budget of the nitrogen cycle at the main station in the Gotland Sea is give

Beschreibung: Somatic growth and nucleic acid content were studied in North Sea houting Coregonus oxyrhinchus larvae fed exclusively on dry diets at 2 temperatures (8.4 and 17.5°C) during a 32 d rearing experiment. The higher temperature enhanced growth significantly. Mean dry weights at the end of the experiment were 3.6 mg (SD = 1.07, range 1.4 to 5.7 mg) and 31.5 mg (SD = 21.9, range 3.6 to 96.0 mg), and mean standard lengths were 17.7 ± mm (SD = 1.6, range 11.8 to 19.6 mm) and 25.5 mm (SD = 4.2, range 17.0 to 35 mm), respectively. Significant responses to temperature were also found in the nucleic acids. However, these differences were not as remarkable and gave indications that differences in protein growth between treatments was based on protein biosynthesis being driven by the activity of the ribosomes, rather than their number. The use of the degree-day approach to normalize the data clearly showed the temperature-dependence of somatic growth. Only small differences in growth and nucleic acid content were observed in the comparable range of the first 280 degree-days in fed fish. In non-fed fish, the starving potential was very similar (approx. 350 degree-days). Shifts in growth pattern from predominantly hyperplasia to predominantly hypertrophy were detected, with ongoing growth at both temperatures.

Beschreibung: The combined and interactive effects of climatic and ecological factors are rarely considered in marine communities. We designed a factorial field experiment to analyze (1) the interactive effects of ambient UV radiation and consumers; and (2) the effects of photosynthetically active radiation (PAR 400 to 700 nm), UVA (320 to 400 nm) and UVB (280 to 320 nm) radiation on a marine hard-bottom community in Nova Scotia, NW Atlantic. Species recruitment and succession on ceramic tiles were followed for 5 mo. We found strong negative UV effects on biomass and cover of the early colonizing macroalga Pilayella littoralis, whereas UVB was more harmful than UVA radiation. Consumers, mainly gammarid amphipods, increased P. littoralis biomass when UV was excluded, probably through fertilization. These initially strong and interacting UV and consumer effects on total biomass and cover diminished as species succession progressed. Species diversity was not affected by experimental treatments, but significant shifts in species composition occurred, especially at the recruitment stage. Red algae were most inhibited by UV, whereas sedentary invertebrates and some brown algae tended to increase under UV exposure. Consumers suppressed green and filamentous brown algae, but favored the other groups. Again, these effects diminished during the later stages of succession. We conclude that UV radiation can be a significant structuring force in early successional benthic communities, and that consumers can mediate its effects.

Beschreibung: Small cetaceans are susceptible to incidental mortality in the various forms of gillnet fisheries throughout their range. Research conducted since 1994 has shown that acoustic alarms (pingers) emitting high-frequency pulsed sounds effectively reduce the number of harbor porpoise Phocoena phocoena casualties in sink gillnets. However, the mechanisms behind the effects of pingers were still not understood. Until now, advantages and risks associated with their widespread use could not be evaluated. Here we present the results of 2 field experiments: (1) theodolite-tracking of harbor porpoises exposed to a single PICE-pinger in Clayoquot Sound, Vancouver Island, Canada and (2) herring Clupea harengus capture rates in surface gillnets equipped with and without acoustic alarms (Dukane Netmark 1000, Lien, PICE) in the Baltic Sea herring fishery at Rügen Island, Germany. Our results show that harbor porpoises do not seem to react to an experimental net in their foraging area (n = 172 groups, median group size = 2 porpoises). Porpoise distance from the mid-point of the net was distributed around a median of only 150 m (range 4 to 987 m). A net equipped with an acoustic alarm, however, was avoided (n = 44 groups) within audible range (distance distribution median = 530 m, range 130 to 1140 m). The porpoises were thus effectively excluded from the ensonified area. Herring, one of the main prey species of harbor porpoises, were not affected by the acoustic alarms tested (n = 25407 fish captured). The advantages and risks of using acoustic alarms to mitigate by-catch are discussed.

Beschreibung: Pelagic and demersal juvenile Baltic cod Gadus morhua L. were collected on the slope and the top of Rønne bank in the Baltic Sea during 2 cruises in November and December 1998. The objective of this study was to evaluate distinct changes in otolith increment width observed in demersal juveniles by comparison with laboratory-reared individuals, and to investigate the factors determining variation in these increments. The different increment-width patterns were identified with a method based on the widths of consecutive increments. Otolith increment widths of juvenile cod were found to be highly variable within and between individuals, in both the experimental and the field samples. The first change in increment pattern observed in the field samples was related to settling. The formation periodicity of increments within the different pattern intervals was confirmed with a growth model based on otolith growth rates of juvenile cod reared in the laboratory under different conditions. In this model, otolith growth rate was expressed as a function of rearing temperature and fish dry weight. Otolith growth of the field samples was calculated using ambient temperatures obtained from a 3D-circulation model. The best fit to observed otolith growth rates was obtained under the assumption that fish on the slope performed daily vertical migrations between the warm surface layer and the cold bottom layer. The data suggested that fish stayed in the surface layer during the first increment-pattern interval, performed vertical migrations during the second interval, and stayed in association with the seafloor in the subsequent interval, corresponding to the time after the breakdown of the thermocline.

Beschreibung: The role of transparent exopolymer particles (TEP) and dissolved organic carbon (DOC) for organic carbon partitioning under different CO2 conditions was examined during a mesocosm experiment with the coccolithophorid Emiliania huxleyi. We designed 9 outdoor enclosures (similar to11 m(3)) to simulate CO2 concentrations of estimated 'Year 2100' (similar to710 ppm CO2), 'present' (similar to410 ppm CO2) and 'glacial' (similar to190 ppm CO2) environments, and fertilized these with nitrate and phosphate to favor bloom development. Our results showed fundamentally different TEP and DOC dynamics during the bloom. In all mesocosms, TEP concentration increased after nutrient exhaustion and accumulated steadily until the end of the study. TEP concentration was closely related to the abundance of E. huxleyi and accounted for an increase in POC concentration of 35 2 % after the onset of nutrient limitation. The production of TEP normalized to the cell Abundance of E. huxleyi was highest in the Year 2100 treatment. In contrast, DOC concentration exhibited considerable short-term fluctuations throughout the study. In all mesocosms, DOC was neither related to the abundance of E. huxleyi nor to TEP concentration. A statistically significant effect of the CO2 treatment on DOC concentration was not determined. However, during the course of the bloom, DOC concentration increased in 2 of the 3 Year 2100 mesocosms and in 1 of the present mesocosms, but in none of the glacial mesocosms. It is suggested that the observed differences between TEP and DOC were determined by their different bioavailability and that a rapid response of the microbial food web may have obscured CO2 effects on DOC production by autotrophic cells.

Beschreibung: The effect of methane released from decomposing surficial gas hydrates (SGH) on standing stocks and activities of the small-sized benthic biota (SSBB; i.e. bacteria, fungi, protozoa, and meiobenthic organisms) was studied at about 790 m water depth, at the Hydrate Ridge, Cascadia subduction zone. Presence of SGH and elevated sulfide concentrations in the sediment were indicated by extensive bacterial mats of Beggiatoa sp. and clam fields of the bivalve mollusc Calyptogena sp. Vertical and horizontal distribution patterns of the SSBB biomass were derived from DNA and total adenylate (TA) sediment assays. Potential bacterial exoenzymatic hydrolytic activity was measured using fluorescein-di-acetate (FDA) as substrate. Estimates of chemoautotrophic production of particulate organic carbon (POC.) were determined by 14CO2 uptake incubations. Inventories of chl a and pheopigments were determined as parameters of surface water primary produced POC input. Average SSBB biomass in clam field sediments integrated over the upper 10 cm (765.2 gC m-2, SD 190.1) was 3.6 times higher than in the adjacent control sites (213 gC m-2, SD 125). Average SSBB biomass in bacterial mat sediments, which were almost devoid of eukaryotic organisms 〉 31 µm, was 209 gC m-2 (SD 65). Significant correlations between FDA, DNA and plant pigments imply that productivity of the SSBB at SGH sites is only partially uncoupled from the primary production of the surface water. Areal estimates of autotrophic Corg production at control sites, bacterial mat sites and in clam field sites were 5.7, 59.7 and 190.0 mgC m-2 d-1, respectively. Based on different models predicting vertical POC fluxes from surface water primary production and water depth, these autotrophic POC productions account for 5 to 17% (controls), 35 to 68% (bacterial mats), and 63 to 87% (clam fields) of the bulk POC (sum of allochthonous POC input through the water column and sedimentary autochthonous autotrophic POC production) provided at the various sites. At SGH sites inventories of chl a and pheopigments, integrated over the upper 10 cm of the sediment, were half of that found at the control sites. This might be due to enhanced degradation of phytodetritally associated organic matter. The resulting low molecular weight organic carbon compounds might stimulate and fuel sulfate reduction, which is conducted in a microbial consortium with anaerobic methane consuming archaea. This syntrophic consortium might represent a prominent interface between gas hydrate derived carbon and allochthonous Corg flow. We infer that degradation kinetics of SGH is affected by, e.g., seasonally varying input of allochthonous organic matter.

Beschreibung: This study tested whether the extreme scarcity of larger nanophytoplankton and microphytoplankton in the Gulf of Aqaba and in the open northern Red Sea is caused by nutrient limitation or by selective removal by grazers. Samples of near surface phytoplankton were incubated on board under a fully factorial combination of release from grazing pressure and release from nutrient stress. Release from grazing pressure by different size classes was obtained by sieving through 100, 20, and 10 µm size mesh screens. Release from nutrient stress was obtained by enrichment of Si alone and a full enrichment by N, P, Si and trace elements. Growth rates of most phytoplankton taxa showed a strong, positive response to the full nutrient enrichment and a weaker, but significant response to grazer exclusion. Several diatom taxa showed a weak positive response to Si enrichment. Thus, bottom-up control of medium-sized algae appears to be more important than top-down control.

Beschreibung: Chromophoric dissolved organic matter (CDOM) represents the optically active fraction of the bulk dissolved organic matter (DOM) pool. Recent evidence pointed towards a microbial source of CDOM in the aquatic environment and led to the proposal that phytoplankton is not a direct source of CDOM, but that heterotrophic bacteria, through reprocessing of DOM of algal origin, are an important source of CDOM. In a recent experiment designed at looking at the effects of elevated pCO2 on blooms of the coccolithophorid alga Emiliania huxleyi, we found that despite the 3 different pCO2 levels tested (190, 414 and 714 ppm), no differences were observed in accumulation of CDOM over the 20 d of incubation. Unlike previous mesocosm experiments where relationships between CDOM accumulation and bacterial abundance have been observed, none was observed here. These results provide some new insights into the apparent lack of effect of pCO2 on CDOM accumulation in surface waters, and question the previously proposed mechanisms and rates of CDOM production in natural phytoplankton blooms.

Beschreibung: Increased nutrient loading favors macroalgal blooms in eutrophied coastal ecosystems. The main counteracting factor on this bottom-up support is top-down control by consumers. We asked (1) whether herbivore control on 2 bloom-forming macroalgae in the Baltic Sea varies between different algal life stages, (2) whether herbivores selectively feed on Enteromorpha spp. (Chlorophyceae) thereby supporting dominance of Pilayella littoralis (Phaeophyceae), and (3) whether various herbivore species differ in their effects. In comparative field and laboratory experiments, we analyzed herbivore pressure and selectivity on germling density and adult thalli of Enteromorpha spp. and P. littoralis. In the field, herbivores reduced macroalgal recruitment by 80% within 14 d indicating strong herbivore control at early life stages. Recruits of Enteromorpha spp. were significantly preferred over P. littoralis Adult thalli of both algae showed similar growth rates, but grazing rates were significantly higher on Enteromorpha spp. In laboratory experiments, Idotea chelipes (Isopoda), Littorina saxatilis (Gastropoda) and Gammarus locusta (Amphipoda) strongly reduced germling density, whereas effects of L. littorea were weak. I. chelipes and L. saxatilis significantly preferred germlings of Enteromorpha spp. over P. littoralis. I. chelipes had strong effects on adult Enteromorpha spp. but not on P. littoralis. The effects of G. locusta and L. littorea on both adult algae were smaller and not selective, and L. saxatilis did not feed on adults at all. Different herbivore feeding modes are discussed. We conclude that strong and selective herbivory on juvenile and adult Enteromorpha spp. favors dominance of P. littoralis in the Baltic. An effective control of macroalgal blooms in eutrophied areas can be supported by sustaining high herbivore density and species richness considering the variable and complementary effects of different herbivore species on different algal life stages.

Beschreibung: Experiments were carried out on Georges Bank, a productive coastal region in the northwestern sector of the North Atlantic Ocean, and in the oligotrophic western Sargasso Sea to examine the effects of nutrient (inorganic nitrogen and phosphorus) and organic carbon (glucose) additions on bacterial and phytoplankton growth. Four experiments were conducted in each environment. Phytoplankton growth was monitored over a 36 h period by following changes in the concentration of chlorophyll in unfiltered seawater and in seawater prefiltered through 5 μm screening to reduce grazing pressure. Bacterial production was estimated initially and after 24 h using the 3H-thymidine (TdR) method in unfiltered seawater and in 1 μm filtrate. Phytoplankton biomass increased significantly in response to nutrient additions in all but 1 experiment, whereas chlorophyll concentrations remained unchanged or decreased in all of the unamended (control) treatments or treatments supplemented with glucose. Responses of the phytoplankton community were similar for the 〈5 μm and unfiltered treatments. Bacterial production increased after 24 h in all of the treatments on Georges Bank, and there was little effect of nutrient or glucose addition in unfiltered seawater relative to unamended controls. However, glucose addition to the 〈1 μm filtrate caused substantial increases in bacterial production relative to controls and N/P-amended treatments in 2 of the experiments from this environment. Glucose had no stimulatory effect (relative to unamended treatments) in 3 of the 4 Sargasso Sea experiments, and only a marginal effect in the fourth. However, the addition of inorganic nitrogen and phosphorus in the latter ecosystem resulted in higher bacterial production (relative to unamended treatments or glucose addition) in 2 of the experiments with unfiltered seawater, and very large increases in 3 of the experiments with 1 μm filtrate. The magnitude of the changes in bacterial production differed greatly between unfiltered and filtered seawater in both ecosystems, indicating an important role for bacterial grazers in controlling bacterial population growth. The results of this study indicate different nutritional restraints on bacterial production in these contrasting environments.

Beschreibung: As one of the elements for a model on the food requirements of Humboldt penguins Spheniscus humboldti we determined, via gas respirometry, metabolic rates while swimming and resting in water. During rest in water at 19°C Humboldt penguins (mean body mass 3.6 kg) required 5.95 W kg-1. This corresponds to a thermal conductance in water of 0.2975 W (kg °C)-1 (at Ta 19°C and assuming a Tb of 39°C). When swimming in a 20 m long channel, metabolism rose from 8 W kg-1 at a speed of 0.6 ms-1 to 23.1 W kg-1 at 2.2 m s-1. Transport costs (the cost to move 1 kg of body mass over a distance of 1 m) reached a minimum at 1.4 ms-1 with 8.1 J (kg m)-1, which corresponds to 0.89 J (Nm)-1. We corrected for acceleration and deceleration in the channel to determine transport costs of free-ranging Humboldt penguins travelling at sea, which were calculated as 7 J (kg m)-1 (0.71 J [Nm]-1), at 1.7 m s-1. Birds feeding chicks need to balance the costs of either (1) returning to the breeding island for the night and travelling back to the feeding grounds in the morning or (2) incurring increased thermoregulatory costs associated with resting at sea overnight. Simple calculations show that at water temperatures of 19°C we expect Humboldt penguins to show a tendency to remain at sea overnight if foraging areas are 〉4 km from their island. In colder waters (12°C), this distance increases to 〉9 km. Using previously published data on at-sea activity of Humboldt penguins, we found that foraging costs during chick rearing amount to 340 g anchovies d-1. Finally, we present a general model to convert Humboldt penguin activity data at sea to food requirements.

Beschreibung: The large temporal and spatial variability in carbon isotope fractionation of marine phytoplankton (ε p) is thought to reflect differences in environmental conditions. Meaningful interpretation of this variability requires an understanding of the processes responsible for phytoplankton isotope fractionation. While numerous factors have been suggested to potentially influence ε p, recent theoretical and experimental evidence has emphasized the primary role of phytoplankton growth rate (µ) and CO2 concentration ([CO2aq]) in controlling ε p. Experimental examination of the relationship of ε p with µ and [CO2aq] in studies using different experimental approaches, however, has yielded inconsistent results. Here we directly compare new and previously published data on ε p as a function of CO2 concentration and growth rate for the marine diatom Phaeodactylum tricornutum. When grown under nitrogen-deficient conditions (nitrate-limited chemostat), ε p of P. tricornutum decreases with increasing growth rate. In contrast, under N-replete conditions ε p values are considerably lower at comparable growth rates and CO2 concentrations and are largely insensitive to a 3-fold increase in growth rate due to increasing photon flux density. In both experimental approaches, ε p shows a relatively small CO2 sensitivity in the range of CO2 concentrations naturally occurring in the ocean (8 to 25 µmol kg-1). Below ca 5 µmol CO2 kg-1, a strong decline in ε p with decreasing [CO2aq] is observed. The apparent difference in ε p responses between nitrate-limited and light-controlled cultures of P. tricornutum suggests a principal difference in carbon acquisition for different growth-rate-limiting resources. A mechanistic explanation is proposed and potential implications for the interpretation of phytoplankton carbon isotope fractionation are discussed.

Beschreibung: The sediment beneath and at various distances from the fish farm cages in Cephalonia bay (Eastern Mediterranean) was investigated seasonally through sediment profiling imagery (SPI) as well as through monitoring of geochemical variables and macrofaunal assemblages. The SPI images (SPI) repeatedly showed the same qualitative pattern along the benthic enrichment gradient with readily identifiable attributes such as depth of dark sediment, signs of outgassing and bioturbation marks. Quantitative comparisons showed that a large number of SPI attributes showed significant positive or negative correlation with geochemical and biological attributes describing the effect of fish farming on the seabed. All multivariate patterns obtained through the analysis of SPI attributes were highly correlated to those obtained from standard multivariate analysis of macrofauna during the respective seasons. It is argued that SPI provides an integrated assessment of the sedimentary conditions and therefore may be used as a complement of or even a substitute for standard sampling methods when mapping the effects of aquaculture on silty substrates

Beschreibung: At Hydrate Ridge (HR), Cascadia convergent margin, surface sediments contain massive gas hydrates formed from methane that ascends together with fluids along faults from deeper reservoirs. Anaerobic oxidation of methane (AOM), mediated by a microbial consortium of archaea and sulfate-reducing bacteria, generates high concentrations of hydrogen sulfide in the surface sediments. The production of sulfide supports chemosynthetic communities that gain energy from sulfide oxidation. Depending on fluid flow, the surface communities are dominated either by the filamentous sulfur bacteria Beggiatoa (high advective flow), the clam Calyptogena (low advective flow), or the bivalve Acharax (diffusive flow). We analyzed surface sediments (0 to 10 cm) populated by chemosynthetic communities for AOM, sulfate reduction (SR) and the distribution of the microbial consortium mediating AOM. Highest AOM rates were found at the Beggiatoa field with an average rate of 99 mmol m-2 d-1 integrated over 0 to 10 cm. These rates are among the highest AOM rates ever observed in methane-bearing marine sediments. At the Calyptogena field, AOM rates were lower (56 mmol m-2 d-1). At the Acharax field, methane oxidation was extremely low (2.1 mmol m-2 d-1) and was probably due to aerobic methane oxidation. SR was fueled largely by methane at flow-impacted sites, but exceeded AOM in some cases, most likely due to sediment heterogeneity. At the Acharax field, SR was decoupled from methane oxidation and showed low activity. Aggregates of the AOM consortium were abundant at the fluid-impacted sites (between 5.1 × 1012 and 7.9 × 1012 aggregates m-2) but showed low numbers at the Acharax field (0.4 × 1012 aggregates m-2). A transport-reaction model was applied to estimate AOM at Beggiatoa fields. The model agreed with the measured depth integrated AOM rates and the vertical distribution. AOM represents an important methane sink in the surface sediments of HR, consuming between 50 and 100% of the methane transported by advection.

Beschreibung: Sediment incubation microcosms, multitrap apparatus and water column variables have been employed to describe the dynamic changes in benthic-pelagic coupling between nutrient pools in the Pontevedra ria, NW Spain, during spring and summer 1998. A comparison of the chemical characteristics of suspended and bed sediment together with sediment nutrient effluxes revealed that particulate organic nitrogen and carbon were progressively depleted upon transit through the nutrient pools. The main fate of particulate organic nitrogen reaching the bed sediment is denitrification, although resuspension is also important. An estimate of net denitrification (Dnet) was made at 2 muddy sites in the ria with a mass balance at the benthic boundary layer. First-order approximations calculate Dnet to be 178 and 182 µmolN m-2 h-1 at the 2 stations, and agree well with previous modelling estimates. Denitrification is highest when upwelling relaxes and the flux of organic matter to the sediment increases. Regular inputs of offshore water ensure water renewal and re-oxygenation of bottom waters, thus preventing anoxia, particularly in the summer. With upwelling, large quantities of ammonium are effluxed to the water column (250 µmolNH4+ m-2 h-1), probably as a result of the bed sediment resuspension engendered by upwelling and stirring of phytodetrital fluff held in suspension as neutrally buoyant material above the sediment surface. We hypothesise that hydrodynamical processes play an important role in determining the quantity of nutrients remineralised in the Pontevedra ria and, in the case of nitrogen, the rate of denitrification at the benthic boundary layer.

Beschreibung: The carbon and nitrogen content of transparent exopolymer particles (TEP) was determined and related to the concentration of TEP as quantified by a colorimetrical method. TEP were produced in the laboratory from dissolved precursors by laminar or turbulent shear. Dissolved precursors were obtained by 0.2 µm filtration from diatom cultures, with or without nutrient reduction, and from natural diatom populations. The relationship between carbon and TEP was significant, linear and species-specific. Carbon concentration of TEP derived from this relationship concurred with previous findings. Shortage of silicic acid or nitrate in the culture media had no effect on the carbon content of TEP. Molar C:N ratios of TEP were above the Redfield ratio, with a mean value of 26. It is suggested that the nitrogen fraction of TEP can be explained by adsorption of dissolved organic nitrogen (DON) onto TEP. Based on the newly established relationship, concentrations of TEP-derived carbon (TEP-C) were calculated for the Baltic Sea, the coastal Pacific, the North East Atlantic and the Northern Adriatic Sea.

Beschreibung: In the Baltic Sea, rye tested how short nutrient pulses of different lengths and frequencies affect macroalgae, epiphytes, grazers and their interactions. We hypothesized that even small-scale variations in nutrient supply may have significant impacts by favoring fast-growing epiphytes which can cause large-scale declines of canopy-forming macroalgae. In a factorial field experiment single plants of the canopy-forming macroalga Fucus vesiculosus with and without epiphytes were exposed to pulses of elevated nutrients (N and P) over 25 d. Five 1 h pulses given every 5 d had no significant effects. A single 5 h pulse increased the epiphyte load but not F. vesiculosus growth rate. In contrast, increasing epiphyte load caused F. vesiculosus growth rate to decline and attracted higher densities of gastropod grazers. These results indicate that a single nutrient pulse can have rapid direct and indirect effects on macroalgae and their associated epiphytes and grazers. Temporal variability of nutrient supply (five 1 h vs one 5 h pulse) plays a significant role in determining the response of primary producers and consumers to elevated nutrients.

Beschreibung: An unusually large number of replicated macrofaunal samples (70) was taken from the Western Baltic in May 1995 for a ringtest in an ICES/HELCOM intercalibration exercise. This data set was employed in this study in order to investigate the performance of numerical methods used for predicting species richness and to assess the accuracy of the estimates of abundance and diversity currently used in benthic ecology. The results of this study indicate that: (1) more than 10 replicates are required in order to include in the data set more than two-thirds of the species found in 70 replicates, and more than 53 replicates are required in order to include 95% of the species; (2) estimates of average abundance and of average Shannon-Wiener diversity index using 5 replicates could result in less than 40% error; this could be less than 30% for 10 replicates and less than 5% for 70 replicates; (3) both types of species-richness predictions (jackknife estimate and S 0) increased with increasing number of samples used in the calculations, indicating that their ability to assess overall species richness in the community is rather limited; in particular, it is shown that jackknife overestimates and S0 slightly underestimates species richness. Different configurations of the S0 method were tested in order to optimize its performance, and it was found that both truncation and increasing sampling lag result in increased and stabilized estimates of species richness.

Beschreibung: Gas hydrates occur at the sediment surface on the southern summit of Hydrate Ridge, Cascadia convergent margin. The hydrates are found in mounds several meters in diameter and up to 2 m high, and are covered by sediment and mats of the filamentous sulfur-oxidizing bacteria Beggiatoa. The mounds are surrounded by vesicomyid clams (Calyptogena pacifica, C. kilmeri), which in turn are encircled by solemyid bivalves (Acharax sp.). The zonation pattern of 3 species (Calyptogena spp. and Acharax sp., which harbor chemoautotrophic bacteria in their gills, and the chemoautotrophic Beggiatoa), is also reflected in a change in the entire community structure. Beggiatoa, Calyptogena spp. and Acharax sp. are shown to be characteristic species for the different communities. The Beggiatoa community directly overlaying the gas hydrates consists of seep endemic species in high densities: gastropods (Provanna laevis, P. lomana, Pyropelta corymba, Hyalogyrina sp. nov.), bivalves (Nuculana sp. nov.) and polychaetes (Ampharetidae, Polynoidae, Dorvilleidae). Based on pooled samples, the rarefaction curves show a decrease in species diversity in the Beggiatoa and Calyptogena communities. The hydrogen sulfide gradients in the porewater of sediments below the different communities dominated by either Beggiatoa, Calyptogena spp. or Acharax sp. vary by 3 orders of magnitude. The diffusive sulfide flux based on the measured sulfide concentration gradients is highest in Beggiatoa sp. communities (23 ± 13 mol m-2 yr-1), slightly less in Calyptogena communities (6.6 ± 2.4 mol m-2 yr-1), and low in Acharax communities (0.05 ± 0.05 mol m-2 yr-1). The difference in the sulfide environment is a factor influencing the distribution patterns of the chemoautotrophy-dependant and heterotrophic species at the deep-sea sediments containing gas hydrate.

Beschreibung: A colony of Humboldt penguins Spheniscus humboldti in central Chile was monitored from August 1995 to July 2000 to determine patterns of breeding and colony attendance and how these were affected by climatic (rainfall) and oceanographic (El Niño) factors. Nests were periodically checked for contents and roosting birds were counted from vantage points. Two main breeding events were observed: between August and January (spring event) and between April and June (autumn event). Whereas the spring event regularly produced offspring, the autumn event was systematically affected by rains, causing considerable nest desertion. Adults were present in the colony from August to May, abandoning the colony during winter after the nests were flooded. Juveniles occurred only between November and March. Adults moulted mainly in February, while juveniles moulted in January. During the 1997/98 El Niño episode, the number of breeding pairs was 55 to 85% lower than the mean, the onset of nesting was delayed, and abnormally heavy rainfall flooded nests. While the number of breeding pairs was significantly related to sea surface temperature anomalies (SSTA), breeding success was not. The attendance of adults and juveniles at the colony during El Niño was 25 and 73% lower, respectively, than the mean attendance. This 2-peak breeding strategy of Humboldt penguins appears to have evolved in response to the more favourable oceanographic and climatic conditions of Perú, where breeding is continuous and not interrupted by rains. Although less productive, the species probably maintains its autumnal breeding in central Chile because this provides additional offspring to supplement those regularly produced during the spring event.

Beschreibung: Generalisations on the combined effects of consumers and resources on autotrophs in aquatic food webs largely rely on freshwater studies. In this study, we tested these general concepts with marine benthic microalgae, which are important components of coastal food webs. We manipulated nitrogen availability and herbivore presence in a factorial field experiment in the Western Baltic Sea. Moreover, we investigated how herbivore control varied among 3 sites and 2 seasons and tested for trophic cascades by enhancing demersal fish density at 2 sites. Nitrogen availability and herbivore presence had strong and antagonistic effects on microalgal biomass, species composition and diversity. Herbivores significantly reduced algal biomass, whereas nutrient enrichment led to an increase in biomass. Herbivore effects on microalgal biomass increased with increasing nitrogen availability, indicating a functional response of herbivores to nutrient enrichment. The response of microalgae at the species level suggested a trade-off between nutrient use and grazing resistance which appeared to be linked to algal growth form. Compared to other growth forms, large erect species were most responsive to both nitrogen loading and herbivory. Grazing reduced microalgal diversity at low nutrient supply, but enhanced it at high nutrient supply. Herbivore effects varied considerably among different sites and were stronger in spring than in summer. Manipulations of fish density during summer did not have any effects on microalgal community structure. In conclusion, our results demonstrate that herbivores and nutrients have strong and balancing effects on marine microbenthic community structure.

Beschreibung: Zooplankton grazing on bacterio- and phytoplankton was studied in the Gulf of Aqaba and the Northern Red Sea during Meteor Cruise Me 44-2 in February-March 1999. Protozoan grazing on bacterioplankton and autotrophic ultraplankton was studied by the Landry dilution method. Microzooplankton grazing on phytoplankton 〉6 µm was studied by incubation experiments in the presence and absence of microzooplankton. Mesozooplankton grazing was studied by measuring per capita clearance rates of individual zooplankton with radioactively labelled food organisms and estimating in situ rates from abundance values. Protozoan grazing rates on heterotrophic bacteria and on algae 〈6 µm were high (bacteria: 0.7 to 1.1 d-1, ultraphytoplankton: 0.7 to 1.3 d-1), while grazing rates on Synechococcus spp. were surprisingly low and undetectable in some experiments. Mesozooplankton grazing was weak, cumulative grazing rates being ca. 2 orders of magnitude smaller than the grazing rates by protozoans. Among mesozooplankton, appendicularians specialised on smaller food items and calanoid copepods on larger ones.

Beschreibung: Although the 'intermediate disturbance hypothesis' (IDH) was originally postulated for marine hard-bottom communities, it has rarely been tested for this community type. We experimentally examined the effects of ambient plus artificially enhanced UV-B radiation (eUV-BR) along an intensity gradient on the abundance of individual species, species composition, and diversity of a macrobenthic community in the western Baltic, Germany. Plots were either exposed each day for 0.5, 1, 2, 4, and 8 h to the maximum expected future UV-BR or were left untreated (= ambient irradiance). Species recruitment and succession on artificial substrata were followed for 5 mo. Transient unimodal patterns for species richness and diversity H' were produced along the UV-BR gradient early in succession. The density of the competitively superior mussel Mytilus edulis and the abundance of the foliose green alga Ulvopsis grevillei were positively related to UV-BR exposure; suggesting conditions different from those predicted by the IDH-generated unimodal patterns. Protective shading of substratum by U. grevillei and the resultant mitigation of the UV-BR impact for succeeding colonizers may be important in this context. Species composition differed persistently among eUV-BR treatments. In contrast to mussels and U. grevillei, the abundance of red algae was adversely affected by eUV-BR along the disturbance gradient. Our results suggest that expected future UV-BR levels will have limited influence on the diversity of shallow-water macrobenthic fouling assemblages in the western Baltic, whereas their species composition may be affected over longer periods.

Beschreibung: Dilution experiments were used to examine growth and grazing mortality rates among Prochlorococcus and Synechococcus populations in the Sargasso Sea and California Current. In these experiments, deviation from linearity in the relationship between dilution and net growth rate was significant in a large number of cases. An alternative, more conservative approach for estimating growth and grazing mortality rates (independent of the shape of this relationship) was therefore employed. Growth rates estimated by this approach ranged from 0.32 to 0.76 and 0.37 to 0.67 d-1 for Prochlorococcus and Synechococcus, respectively. Grazing mortality rates ranged from 0.25 to 0.85 and 0.13 to 0.51 d-1, respectively. Cell-cycle-based growth rate estimates were consistent with these dilution-based rates. Nutrient amendment had little affect on picocyanobacterial growth rates, but did stimulate grazing mortality (and in some cases changed the apparent functional response of the grazer community) in a number of experiments. We hypothesize that improved food quality in nutrient-replete picoplankton cells may be responsible for these changes. Diel patterns of picocyanobacterial abundance in the Sargasso Sea experiments suggest that grazing activity varied strongly over the diel cycle, with low grazing activity during the first half of the light period. Growth rate and abundance were not positively correlated among or within picocyanobacterial groups, as might be expected if physiologically mediated controls were the dominant forces regulating these populations.

Beschreibung: Factors responsible for aggregations of jumbo flying squid Dosidicus gigas, an important component of the marine food web and target of commercial fisheries off the Costa Rica Dome in the Eastern Tropical Pacific Ocean (ETP), were examined during 2 years of different extreme oceanographic conditions: fall 1997 El Niño and fall 1999 La Niña. A high abundance of squid occurred in association with the well-developed countercurrent ridge (upwelling) off the Costa Rica Dome during fall 1997, but not during fall 1999, when the countercurrent ridge was less developed. Two features of the well-developed countercurrent ridge were considered important for the occurrence of high jumbo flying squid concentrations. Firstly, subsurface chlorophyll a (chl a) maxima were formed along the countercurrent ridge, resulting in integrated chl a concentrations in the upper 100 m being relatively high considering the generally low productivity of the ETP during an El Niño event. Secondly, a strong salinity front formed along the North Equatorial Countercurrent, which is possibly responsible for retention of jumbo flying squid in the ridge. Large yellowfin tuna Thunnus albacares, which mainly feed on micronekton (small fishes, cephalopods and swimming crabs), as do jumbo flying squid, were also more highly concentrated along the countercurrent ridge during 1997 than during 1999. It was noted that skipjack tuna Katsuwonus pelamis and small yellowfin tuna, which mainly feed on zooplankton, were associated with the equatorial ridge in the ETP, indicating that prey faunal components may also play an important part in the close association of jumbo flying squid with the countercurrent ridge.

Beschreibung: Squids typically demonstrate considerable plasticity in individual growth rates. However, it is not known if individuals growing at different rates also differ at lower levels of organisation. We wished to determine if Sepioteuthis lessoniana individuals that were larger than predicted for their age differed in their digestive gland and mantle muscle tissue proximal composition or mantle muscle structure, compared with individuals that were smaller for their age than predicted. The residual, the difference between the observed size-at-age and that predicted by the growth equation, was used as a measure of the difference in an individual's lifetime growth from the population average. Individual squid varied considerably in their size-at-age, with juveniles showing less variation than adults. Juveniles had greater concentrations of lipid in their muscle tissue, perhaps due to an emphasis on storing energy reserves in this critical period of their life. Differences in biochemical constituents in both the digestive gland and muscle tissue were not related to the size-at-age of individuals, despite biochemical make-up being the lowest organisational level of growth. This may be due to whole animal growth and changes in biochemical composition occurring on different time scales. There was no relationship between the size-at-age of individuals and average mantle muscle fibre size. A strong relationship, however, existed between the size of mantle muscle blocks and the size-at-age of individuals for both juvenile and adult individuals, suggesting that larger muscle blocks are related to both body size and faster individual growth rates. This study demonstrates a clear relationship between mantle muscle structure and growth and the size-at-age of S. lessoniana individuals.

Beschreibung: Age, growth and maturity parameters were examined for the southern Australian ommastrephid squid Nototodarus gouldi. Squid were obtained from the fishing ports of: Ulladulla, New South Wales; Port Lincoln, South Australia; Lakes Entrance, Victoria; and Hobart, Tasmania. Squid were collected during 2 seasonal periods: summer/autumn-caught (warm-season squid) and spring-caught (cool-season squid) over 2 consecutive years (2000, 2001). N. gouldi is a sexually dimorphic species, with females generally reaching larger sizes than males. Initial genetic analysis has found only a single species in Australian waters. Statolith ageing revealed that squid completed their life cycle in 〈1 yr, and appear to hatch throughout the year. Trends in size, growth and maturity varied considerably between sites, seasons and years. Squids hatched in summer/autumn grew consistently faster than squid that hatched in winter/spring, presumably due to the influence of temperature on growth. Squid in 1999/2000 also grew faster than squid in 2000/2001. Growth of female squid in winter correlated with sea surface colour (SSC) during peak hatch periods, but the SSC relationship did not exist for males. Ulladulla squid were generally smaller, younger, had smaller gonads than most other squid and were possibly a smaller morph of the species. Tasmania and Lakes Entrance tended to have larger older individuals with larger gonads, while Port Lincoln was variable and intermediate. However, during spring 2001 both Tasmania and Port Lincoln had individuals that were much smaller than those of the other seasons for these sites, and were more like those from Ulladulla. Trends in age of mature individuals showed considerable variability (over 100 d from youngest to oldest) and there appeared to be a cline across all sites and seasons. Arrow squid appear to reveal marked plasticity in age, growth and maturity parameters, but currently the extent to which the environment or genetics control plasticity is unclear.

Beschreibung: We fertilised 12 mesocosms with NE Atlantic phytoplankton with different Si:N ratios (0:1 to 1:1). After 1 wk, we added mesozooplankton, mainly calanoid copepods at natural densities to 10 of the mesocosms; the remaining 2 mesocosms served as controls. A trend of increasing diatom dominance with increasing Si:N ratios and species-specific correlations of diatoms to Si:N ratios were not changed by the addition of mesozooplankton. Large unicellular and chain-forming diatoms, thin-walled dinoflagellates (Gymnodiniales) and ciliates were reduced by copepod grazing while armoured dinoflagellates remained unaffected. Nanoplanktonic flagellates and diatoms profited from the addition of copepods, probably through release from ciliate grazing.

Beschreibung: Multiple-choice feeding experiments were performed with the isopod Idotea granulosa and the amphipod Gammarus locusta as consumers. In a first experiment, 2 different types of tissues of the brown seaweed Fucus vesiculosus and its main macroepiphytes, Ulva lactuca and Elachista fucicola, were offered. I. granulosa rejected apices of F. vesiculosus and preferred E. fucicola, while G. locusta clearly preferred F. vesiculosus tissue, especially the meristematic apices. In a second experiment, F. vesiculosus tissue with and without E. fucicola was offered together. For I. granulosa, the consumption of F. vesiculosus was enhanced by the presence of the epiphyte, while for G. locusta there was no difference in consumed F. vesiculosus mass. G. locusta, however, showed behavioural rejection of E. fucicola, and thus, the epiphyte acted as Œprotective coating¹. We conclude that host (F. vesiculosus) tissue could be Œco-consumed¹ by mesograzers (I. granulosa) that were attracted by the presence of epiphytes and that these epiphytes therefore may have a 2-fold negative effect on the host (i.e. competion for light, nutrients etc. and attraction of consumers). ŒCo-consumption¹ and Œprotective coating¹ add 2 more facets to the very variable and case-dependent interrelationships of mesograzer-epiphyte-host systems; their relevance in nature, however, remains to be demonstrated.

Beschreibung: Herbivory can greatly modify benthic community structure by affecting the distribution of algal species. To deter herbivores, algae have developed several mechanisms, including the induction of chemical and morphological defenses, which may be influenced by nutrient availability. We tested 4 red (Chondrophycus flagellifera, Cryptonemia seminervis, Osmundaria obtusiloba, Pterocladiella capillacea), 4 brown (Dictyota menstrualis, Lobophora variegata, Sargassum vulgare, Stypopodium zonale), and 1 green (Codium decorticatum) algae for inducible defenses following exposure to direct consumption by an amphipod community dominated by Elasmopus brasiliensis. In addition, the effects of water-borne cues from nearby grazed conspecifics and non-grazing consumers on the induction of defenses were examined in C. decorticatum under natural and enhanced (200% natural) nutrient levels. Induction of defense was assessed in choice-feeding assays, using live algae or artificial food containing non-polar extracts of amphipod-exposed (treated) and non-exposed (control) algae. Palatability levels, estimated as the relative difference in wet mass due to consumption in feeding assays between grazer-exposed and control plants, declined significantly in 3 species after the acclimatization period. Tissue from the directly consumed red alga P. capillacea (live alga) was significantly less palatable than tissue from the control plants. Likewise, a significant effect was observed in the brown alga L. variegata. Similar, although not statistically significant, trends were observed in 6 other species. For the green alga C. decorticatum, nutrient enrichment did not affect induction of defenses by herbivores, yet unfertilized plants were more palatable than fertilized conspecifics.

Beschreibung: Connell¹s intermediate disturbance hypothesis (IDH) postulates that diversity peaks at an intermediate level of disturbance frequency or intensity. To test the validity of this concept for the species-poor marine hard-bottom community of the Western Baltic, we chose an in situ experimental approach. Undisturbed fouling communities of 2 different successional stages, 3 and 12 mo old, were submitted to various levels of emersion intensities, defined as time spans of continuous exposure to the air d-1. Disturbance levels ranged from 0.25 h up to 12 h of daily exposure. The study on 3 mo old communities was repeated in 2 subsequent years, 1999 and 2000. Species richness, evenness and diversity (Shannon index) were recorded to measure the effect of intensity treatments on community structure. The IDH was confirmed in the first year, when diversity was found to peak at intermediate disturbances. However, for communities of both successional stages, diversity-disturbance relationships were U-shaped or not significant in the second year. This ambiguous picture basically confirms the validity of the mechanisms proposed by the IDH, but shows that their forcing can be masked by fluctuations in environmental parameters, such as climatic conditions. An extension of the model is proposed, that considers diversity enhancement under extreme conditions due to a disturbance-induced change in community structure. Furthermore we discuss a conceptual linkage of the IDH to the multiple stable-state hypothesis. Finally, we found community stability not to be positively correlated with community age and complexity.

Beschreibung: Some marine mammals may increase their underwater locomotor efficiency by taking down little air for dives and descending passively, although at the point of maximum depth they presumably have to use energy to counteract the downthrust, to stop themselves sinking further. Birds, having considerable quantities of body-associated air, would appear not to have this option. However, measurements of locomotor activity and inspiratory behaviour of free-living, diving penguins has revealed that birds regulated the inspired air volume so that upthrust, primarily derived from depth-related changes in air volume, was minimal and constant at the preferred foraging depth. Although this results in minimized costs of travel, it means total body oxygen stores have to vary with depth, something that helps explain why dive duration is so closely correlated with depth in birds.

Beschreibung: We examined how seabirds might be used to study marine environmental variables, which necessitates knowing location and the value of the variable to be studied. Five systems can potentially be used for determination of location: VHF (Very High Frequency) telemetry, PTT (Platform Terminal Transmitters) telemetry, GLS (Global Location Service) geolocation methods, dead reckoning and GPS (Global Positioning System), each with its own advantages with respect to accuracy, potential number of fixes and size. Temperature and light were used to illustrate potential difficulties in recording environmental variables. Systems currently used on seabirds for measurement of temperature respond slowly to environmental changes; thus, they may not measure sea surface temperature adequately when contact periods with water bodies are too short. Light can be easily measured for light extinction studies, but sensor orientation plays a large role in determining recorded values. Both problems can be corrected. The foraging behaviour of seabirds was also examined in order to identify those features which would be useful for determination of marine environmental variables at a variety of spatial and temporal scales. Area coverage by birds is highly dependent on breeding phase and tends to be concentrated in areas where prey acquisition is particularly enhanced. The identification of these sites may be of particular interest to marine biologists. 'Plungers' and 'divers' are potentially most useful for assessment of variables deeper within the water column, with some divers spending up to 90% of their time sub-surface. Few seabirds exploit the water column deeper than 20 m, although some divers regularly exceed 50 m (primarily penguins and auks), while 2 species dive in excess of 300 m. The wide-ranging behaviour of seabirds coupled, in many instances, with their substantial body size makes them potentially excellent carriers of sophisticated environmental measuring technology; however, the ethical question of how much the well-being of birds can, and should, be compromised by such an approach needs to be carefully considered.

Beschreibung: The phylogenetic relationship of sulphur-oxidising endosymbiotic bacteria from bivalves of the families Vesicomyidae (Calyptogena sp. C1, Calyptogena sp. C3), Solemyidae (Acharax sp.) and Thyasiridae (Conchocele sp.) from cold-seep habitats were determined by 16S rDNA nucleotide sequence analyses. The endosymbiotic bacteria form distinct groups within the gamma-Proteobacteria and are well separated from each other and from free-living sulphur-oxidising bacteria of the genera Beggiatoa, Halothiobacillus and Thiomicrospira. The endosymbiotic bacteria of Acharax sp. from cold seeps off Oregon, Indonesia and Pakistan have sequences highly similar to each other but quite distinct from other thiotrophic endosymbionts. This includes endosymbionts from Solemya spp., to which they are distantly related. Symbiotic bacteria of Conchocele sp. from a cold seep in the Sea of Okhotsk are similar to those of Bathymodiolus thermophilus and related species, as shown by their overall sequence similarity and by signature sequences. The endosymbiotic bacteria of Calyptogena spp. from cold seeps off Oregon and Pakistan are closely related to those of other vesicomyids. Endosymbiont species found off Oregon corresponded to 2 different clusters of Calyptogena spp. symbionts in the same samples. The results corroborate the hypothesis of a monophyletic origin of the symbionts in vesicomyid clams, and support the existence of deeply branching groups in solemyid symbionts and of divergent lines and distribution for thyasirid symbionts. The results also indicate that certain symbiont species cluster according to the depth distribution of their hosts, and that in consequence host species together with their symbionts may have undergone depth-specific adaptation and evolution.

Beschreibung: New technology has allowed inter-dive oxygen-loading behaviour to be assessed in free-living Magellanic penguins Spheniscus magellanicus and shows, via pre-dive hyperventilation and oxygen loading, that birds predict both their upcoming dive performance in terms of duration and depth, and foraging success based on the number of prey they are likely to catch. Thus wild penguins make decisions based on complex environmental parameters and this leads to particularly efficient time use as expected according to optimized foraging.

Beschreibung: We determined key chemical parameters and thiosulfate oxidation in low-temperature hydrothermal fluids from the North Fiji Basin. In addition, the bacterial diversity (with the main emphasis on sulfur-oxidizing bacteria) was investigated. The hydrothermal fluids had low concentrations of sulfide (up to 50.0 µM) and increased counts of both total bacteria and sulfur-oxidizing bacteria compared to ambient seawater. Pure cultures of bacteria were isolated from these fluids on media suited for autotrophic, and potentially heterotrophic, sulfur-oxidizing bacteria. Evidence for the abundance of α- and γ -Proteobacteria was obtained from identification of isolated pure cultures. A large number of 16S rDNA sequences of these groups were retrieved from environmental DNA. Representatives of the Cytophaga-Flavobacterium phylum were found by denaturing gradient gel electrophoresis (DGGE) and 16S rDNA sequence information of DGGE bands, although these bacteria could not be isolated with the media used in this study. Evidence for the presence of chemolithoautotrophic sulfur-oxidizing bacteria was found by analysis of environmental DNA, using 16S rDNA-specific primers of various groups of chemotrophic sulfur bacteria. They were isolated in low numbers compared to chemoheterotrophic and mixotrophic sulfur-oxidizing bacteria. In addition, the formation of tetrathionate as major oxidation product of thiosulfate added to hydrothermal fluid samples and to pure cultures of new isolates indicates the importance of chemoheterotrophic sulfur-oxidizing bacteria within the warm vent waters investigated during this study.

Beschreibung: Albatrosses have among the most remarkable travelling capacities of any extant animal. However, previous studies regarding their movements at sea have mainly focused on breeding birds commuting between the nest site and offshore feeding grounds. In this study, we compare the movement patterns and at-sea activity of breeding and inter-breeding black-browed albatrosses Diomedea melanophris from the Falkland Islands. Data were recorded via global location and activity sensors for 26 incubating birds [during single foraging trips lasting 6.8 d on average) and 6 inter-breeding individuals (during non-stop offshore journeys of 127.5 d on average). Our results showed that foraging black-browed albatrosses utilise vast offshore areas (the average foraging area was 102000 +_ 132 000 km2 by incubating birds and 1552 000 * 970 000 km2 by inter-breeding birds). However, mean forag~ngr ange was similar in both groups (691 * 330 km and 680 t 192 km by incubating and interbreeding birds, respectively) as were their core foraging areas and their at-sea activity patterns. Our results thus indicate that black-browed albatrosses from the Falkland Islands, which represent the largest albatross population world-wide (ca 800 000 individuals), mainly rely on marine resources available within the Patagonian Shelf area. Although this hghly productive continental shelf is the largest of the Southern Hemisphere, rapid development of industrial fisheries currently results in the removal of over 1.4 million tonnes of fish and squid per year in this zone. As our data also show significant spatio-temporal overlap between human and albatross fishing activities within the Patagonian Shelf, we anticipate major detrimental effects on the albatross population in terms of competition for food and additional mortality caused by bird bycatch.

Beschreibung: We report how different zooplankton groups (doliolids, cladocerans and copepods) are able to influence the coastal pelagic food web, including the microbial food web, in waters of the NW Mediterranean. We studied the effect of grazing and of grazing-induced nutrient recycling mediated by different types of zooplankton grazing on a natural phytoplankton community. Experiments were conducted in semicontinuous 2-stage chemostats. The 1st stage vessels contained seawater from Blanes Bay, Spain (NW Mediterranean) including its natural phytoplankton community; the 2nd stage vessels contained the same seawater and copepods, cladocerans or doliolids. At daily intervals we transferred part of the medium from the 2nd to the 1st stage flasks, which contained ungrazed algae and excreted nutrients. In this way, the zooplankton could influence phytoplankton dynamics both by selective grazing and by differential excretion of limiting nutrients. In the 2nd stage flasks grazing changed the algal community composition. Doliolids and cladocerans promoted the growth of large algae and copepods shifted the size spectrum towards small sizes. This effect was transferred to the 1st stage flasks. Doliolids, cladocerans and copepods also affected the microbial food web in different ways. Size-selective grazing led to differences in the nanoplankton concentrations. These in turn affected bacterial concentrations in a trophic cascade. The potential to modify a given algal population increased with increasing selectivity of the grazer.

Beschreibung: Both surface texture and littorinid grazing are known to influence the establishment of many shallow-water benthic hard-bottom communities. However, the effects of these factors and their interactions have not yet been investigated in a quantifiable manner. This investigation aims to assess the interactive effect of both factors in a strictly standardized manner. Natural recruitment by diatoms, the barnacle Balanus improvisus and the tube-building polychaete Polydora sp. was monitored under a 2-factorial treatment: grazing by the periwinkle Littorina littorea (Factor 1; 2 levels: I or no snails per plate) on artificial recruitment plates of different initial surface rugosities (Factor 2; 5 levels: smooth, 0.1, 0.5, 1 and 5 mm rugosity elements), In the absence of grazers, barnacle recruitment decreased with increasing initial rugosity, polychaete recruitment peaked at intermediate rugosities, and diatoms exhibited contrasting recruitment patterns in an in vitro and an in situ experiment. When preferred recruitment sites coincided for Polydora and B. improvisus, a competition for space could be inferred from a negative correlation between the 2 species. However, when the overlap of requirements weakened on the 5 rugosities, the relationship was positive, but was not statistically significant. Grazing efficiency by L. littorea depended on initial rugosity, generally showing minimum values on intermediate rugosities which is attributable to a mismatch between radula dimensions and surface structures in these rugosity classes. Additionally, grazing effects tended to increase with higher prey densities. As all factors-initial rugosity, grazing, colonizer species-interact with each other, the outcome of recruitment under combined factors is difficult to predict from single factor effects.