Methodological challenges in assessing the environmental status of a marine ecosystem: case study of the Baltic Sea

Assessments of the environmental status of marine ecosystems are increasingly needed to inform management decisions and regulate human pressures to meet the objectives of environmental policies. This paper addresses some generic methodological challenges and related uncertainties involved in marine ecosystem assessment, using the central Baltic Sea as a case study. The objectives of good environmental status of the Baltic Sea are largely focusing on biodiversity, eutrophication and hazardous substances. In this paper, we conduct comparative evaluations of the status of these three segments, by applying different methodological approaches. Our analyses indicate that the assessment results are sensitive to a selection of indicators for ecological quality objectives that are affected by a broad spectrum of human activities and natural processes (biodiversity), less so for objectives that are influenced by a relatively narrow array of drivers (eutrophications, hazardous substances). The choice of indicator aggregation rule appeared to be of essential importance for assessment results for all three segments, whereas the hierarchical structure of indicators had only a minor influence. Trend-based assessment was shown to be a useful supplement to reference-based evaluation, being independent of the problems related to defining reference values and indicator aggregation methodologies. Results of this study will help in setting priorities for future efforts to improve environmental assessments in the Baltic Sea and elsewhere, and to ensure the transparency of the assessment procedure.     

Horizontal distribution of photo- and heterotrophic microorganisms on the coastal area of the northern Baltic Sea--a case study

The horizontal distribution of bacteria (BACT), bacterial production(FDC), picoplanktonic cyanobacteria (CYANO), and photosynthetic(PNF) as well as heterotrophic (HNF) nanoflagellates was studiedat the end of June on the SW coast of Finland, the Baltic Sea.Samples were taken at 0 and 10 m depths from three transects500 and 800 m apart. The distance between sampling points was20 m. Cell counts of BACT, CYANO, PNF, HNF and the FDC showed3- to 5-fold deviations in the whole data set. Differences betweenthe duplicate counts within the subsamples were 10–20%.Within the transects, the maximum values were 1.2–3.0times higher than the minimum values. Differences between thetransects were statistically significant for all variables exceptFDC. Deviations in cell numbers between the surface and 10 mwere quite irregular. but often statistically significant. Inmost cases, the organism groups did not correlate significantlywith each other. The horizontal distribution of pelagic microbeswas sufficiently uniform, on a meter scale. for one samplingpoint to give a reliable estimate of microbial densities inthe area under investigation. The differences between the surfaceand the lower stratum of the mixed layer (10 m) were in mostcases statistically significant.     

The broad-scale distribution of five jellyfish species across a temperate coastal environment

Jellyfish (medusae) are sometimes the most noticeable and abundant members of coastal planktonic communities, yet ironically, this high conspicuousness is not reflected in our overall understanding of their spatial distributions across large expanses of water. Here, we set out to elucidate the spatial (and temporal) patterns for five jellyfish species (Phylum Cnidaria, Orders Rhizostomeae and Semaeostomeae) across the Irish & Celtic Seas, an extensive shelf-sea area at Europe’s northwesterly margin encompassing several thousand square kilometers. Data were gathered using two independent methods: (1) surface-counts of jellyfish from ships of opportunity, and (2) regular shoreline surveys for stranding events over three consecutive years. Jellyfish species displayed distinct species-specific distributions, with an apparent segregation of some species. Furthermore, a different species composition was noticeable between the northern and southern parts of the study area. Most importantly, our data suggests that jellyfish distributions broadly reflect the major hydrographic regimes (and associated physical discontinuities) of the study area, with mixed water masses possibly acting as a trophic barrier or non-favourable environment for the successful growth and reproduction of jellyfish species. Handling Editor: K. Martens     

Habitat loss and possible effects on local species richness in a species-poor system: a case study of southern Baltic Sea macrofauna

The effects of habitat loss on local species richness depend on the characteristics of the endangered system (including its total species pool and the distribution of species among the habitats). The present study focuses on the species-poor southern Baltic marine benthic biota. Macrobenthic samples were collected in three habitats: (1) soft bottom covered with vegetation; (2) stony reefs; (3) unvegetated sands. Fourty one percent of 54 observed macrozoobenthic species were habitat specific, while 30% occurred in all three habitats. There were no significant differences in total species richness among the three habitats. The accumulation curves plotted for subsets of data with selected habitats excluded lay below the curve plotted for the whole dataset, but only in one case the 95% confidence intervals of the subset curve did not overlap with those plotted for the whole dataset. The exclusion of samples from selected habitats produced a species richness drop ranging from 9 to 13%. The present study showed that habitat loss in a species-poor area with a relatively large ratio of generalist species cannot produce local species richness declines similar to those predicted for diverse marine systems. However, it must be emphasized that in species-poor systems, the loss of ecological function accompanying habitat loss could be disproportionally higher than that predicted based on decreases in species richness, as some functions are performed by a single species.     

Planktonic ciliated protozoa: their distribution and relationship to environmental variables in a marine coastal ecosystem

The spatial distribution of planktonic ciliated protozoa, physicalvariables and potential food items were measured at northernand southern sites off the west coast of the South Island, NewZealand, to examine which variables were important in structuringthe ciliate community. Ciliates contributed 30% of total zooplanktonbiomass. Ciliate abundance was greatest within 55 km of thecoast on the northern transect, but on the southern transectthe greatest abundance was close inshore and at the stationsoffshore of the continental shelf. The ciliate community wasdominated by small oligotrichs and abundance (114–1139l^–1 was comparable with other oligotrophic marine environments.The depth of the pycnocline had a major influence on the verticaldistribution of ciliates, which were closely correlated withprofiles of phytoplankton biomass and picophytoplankton abundance,but not primary production. The results of this study suggestthat physical factors were more important than biotic factorsin influencing the distribution of protozoan ciliates. The deepmixed layer prevented the formation of deep chlorophyll a orciliate maxima. Few studies of microzooplankton have incorporatedpicophytoplankton distribution, but the importance of smallciliates and picophytoplankton suggests that these groups mayplay a pivotal role in mediating the transfer of organic matterto higher trophic levels in this marine coastal ecosystem.     

Prediction of benthic community structure from environmental variables in a soft-sediment tidal basin (North Sea)

The relationship between benthos data and environmental data in 308 samples collected from the intertidal zone of the H?rnum tidal basin (German Wadden Sea) was analyzed. The environmental variables were current velocity, wave action, emersion time (all of which were obtained from a 2-year simulation with a numerical model) and four sediment grain-size parameters. A grouping of sample stations into five benthos clusters showed a large-scale (>1?km) zoning of benthic assemblages on the tidal flats. The zoning varied with the distance from the shore. Three sample applications were examined to test the predictability of the benthic community structure based on environmental variables. In each application, the dataset was spatially partitioned into a training set and a test set. Predictions of benthic community structure in the test sets were attempted using a multinomial logistic regression model. Applying hydrodynamic predictors, the model performed significantly better than it did when sediment predictors were applied. The accuracy of model predictions, given by Cohen’s kappa, varied between 0.14 and 0.49. The model results were consistent with independently attained evidence of the important role of physical factors in Wadden Sea tidal flat ecology.     

Locomotor activity patterns of nineteen fish and five crustacean species from the Baltic Sea

Synopsis Activity patterns of some common Baltic fish species and macrocrustaceans have been investigated in natural light/dark conditions — in most cases for periods of at least one full year. The fauna of the northern Baltic proper consists of species of both marine and freshwater origin, with both types represented in the study. The different patterns found are discussed in relation to light period, light intensity and to the structure and function of the eyes. The following species were considered nocturnal: Idothea baltica, Mesidothea entomon, Gammarus oceanicus, Palaemon adspersus, Anguilla anguilla, Spinachia spinachia, Pholis gunnellus, Zoarces viviparus, Scopthalmus maximus, and Platichtys flesus. These species were considered diurnal: Clupea harengus, Scardinius erythrophthalmus, Gasterosteus aculeatus, Perca fluviatilis, Gobius niger, Taurulus bubalis, and Cyclopterus lumpus. The next three species showed a crepuscular pattern of activity: Praunus flexuosus, Esox lucius and Gymnocephalus cernus. Four species were considered to show inversions of the activity patterns: Pungitius pungitius, Pomatoschistus minutus, Myoxocephalus scorpius and M. quadricornis.     

Sponge communities of the Antarctic Peninsula: influence of environmental variables on species composition and richness

Sponge communities on the Antarctic continental shelf currently represent one of the most extensive sponge grounds in the world, and all sponge classes are known to occur in the Southern Ocean. Main objectives of this study conducted at the tip of the Antarctic Peninsula were (1) to identify all sampled sponges and (2) to investigate whether the species composition and species richness of Southern Ocean sponge communities in the area of the Antarctic Peninsula are significantly influenced by environmental variables. The studied material originated from 25 AGT catches and was sampled during the expedition ANT-XXIX/3 of RV Polarstern. Samples were collected in three large-scale areas in the vicinity of the Antarctic Peninsula: Bransfield Strait, Drake Passage and Weddell Sea. The following six environmental variables were measured from bottom water samples (except for sea-ice cover): depth (m), light transmission (%), oxygen (µmol/kg), salinity, sea-ice cover (%) and temperature (°C). Two hundred and sixty-three sponge samples were analyzed, and 81 species of 33 genera from all Porifera classes (Calcarea, Demospongiae, Hexactinellida and Homoscleromorpha) were identified. Total numbers of sponge species per sample station ranged from 1 to 29. A detrended correspondence analysis and a backward-stepwise model selection were performed to check whether species composition and richness were significantly influenced by environmental variables. The analyses revealed that none of the measured environmental variables significantly influenced species composition but that species richness was significantly influenced by (1) temperature and (2) the combination of temperature and depth. Results of this study are of crucial importance for development, performance and assessment of future protection strategies in case of ongoing climatic changes at the Antarctic Peninsula.     

Temporal and spatial variation of coastal zooplankton in the Baltic Sea

When analysing temporal variation monthly mean abundances of zooplankton sampled at a eoastal station in the northern Baltic Sea between 1976 and 1988 showed the lowest between-year variation in the summers The coefficients of variation were estimated at 50% for rotifers in June. 70% for cladocerans in August and between 30 and 50% for different copepodite stages in August Moreover in the summers, all dominating zooplankton groups were abundant Estimates of yearly biomass or production of zooplankton must, however, be based on frequent sampling during the whole year because of large differences between months in zooplankton composition and abundance
In 1989. spatial variation was analyzed by sampling zooplankton on three occasions along a 9 8 km transect with 15 stations, 700 m apart All taxonomic/life-stage groupings of zooplankton were heterogeneously distributed Geostatistical analysis did not give a definite answer to what the minimum distance between stations should be to obtain independent samples but indicated that 700 to 1400 m would be sufficient in most cases For most of the zooplankton groupings the coefficients of variation in the sampled area were estimated at 20-40% on all sampling occasions     

Scale‐dependent genetic structure of the Idaho giant salamander (Dicamptodon aterrimus) in stream networks

The network architecture of streams and rivers constrains evolutionary, demographic and ecological processes of freshwater organisms. This consistent architecture also makes stream networks useful for testing general models of population genetic structure and the scaling of gene flow. We examined genetic structure and gene flow in the facultatively paedomorphic Idaho giant salamander, Dicamptodon aterrimus, in stream networks of Idaho and Montana, USA. We used microsatellite data to test population structure models by (i) examining hierarchical partitioning of genetic variation in stream networks; and (ii) testing for genetic isolation by distance along stream corridors vs. overland pathways. Replicated sampling of streams within catchments within three river basins revealed that hierarchical scale had strong effects on genetic structure and gene flow. amova identified significant structure at all hierarchical scales (among streams, among catchments, among basins), but divergence among catchments had the greatest structural influence. Isolation by distance was detected within catchments, and in‐stream distance was a strong predictor of genetic divergence. Patterns of genetic divergence suggest that differentiation among streams within catchments was driven by limited migration, consistent with a stream hierarchy model of population structure. However, there was no evidence of migration among catchments within basins, or among basins, indicating that gene flow only counters the effects of genetic drift at smaller scales (within rather than among catchments). These results show the strong influence of stream networks on population structure and genetic divergence of a salamander, with contrasting effects at different hierarchical scales.     

The influence of cyanobacteria blooms on the attenuation of nitrogen throughputs in a Baltic coastal lagoon

We combined a mass balance approach with measurements of air–water and sediment–water nitrogen (N) exchange to better understand the mechanisms attenuating N throughputs in a eutrophic coastal lagoon. We were particularly interested in how seasonal shifts in external versus internal N fluxes and the transition from diatom- to cyanobacteria- dominated phytoplankton communities influence N storage and loss to the atmosphere. We found that on an annual basis almost all of the N removed by the lagoon was due to sediment storage following the spring diatom bloom. This period was characterized by high riverine inputs of dissolved inorganic nitrogen, high rates of assimilatory conversion to particulate nitrogen (PN), and net accrual of N in sediments. By contrast, the larger summer bloom was associated with low sediment N storage, which we attribute in part to the presence of positively-buoyant cyanobacteria. Low settling rates during cyanobacteria blooms favored export of PN to the Baltic Sea over sediment accrual in the lagoon. In addition, summer dinitrogen (N₂) fixation by cyanobacteria largely offset annual N₂ losses via denitrification. These findings show that cyanobacteria blooms diminish N attenuation within the lagoon by altering the balance of N exchange with the atmosphere and by promoting export of particulate N over sediment burial.     

Long-term changes in the benthic communities of the Pomeranian Bay (Southern Baltic Sea)

Long-term changes in the macrofauna of the Pomeranian Bay were studied by comparing survey data from the 1950s, 1980s, and 1990s. The study area has undergone significant eutrophication during the period of investigation. Biomass of filter-feeding bivalves increased significantly. Spatial distribution patterns of several species have changed. Strong decreases in species richness were caused by oxygen depletion at stations deeper than 15 m.Saduria entomon, Monoporeia affinis, andPontoporeia femorata vanished entirely between 1981 and 1993. Although a causal relationship between simultaneous increases of nutrient levels and macrobenthic biomass cannot be verified, eutrophication is proposed to be the major process affecing changes in macrofauna assemblages. In addition, changes in hydrography and climate increased frequency and severity of oxygen depletion events in the Pomeranian Bay since the mid 1980s.     

Seasonal variations of microphytoplankton assemblages and environmental variables in the coastal zone of Bozcaada Island in the Aegean Sea (NE Mediterranean Sea)

Seasonal changes in the microphytoplankton assemblages were examined in the coastal zone of Bozcaada Island with regard to some major physical and chemical variables. Samples were collected from May 2000 to December 2001 at four stations. A total of 108 dinoflagellates, 102 diatoms, 1 chrysophycean, 3 dictyochophycean, and 1 prasinophycean species were identified and quantified during the study period. Diatoms and dinoflagellates were the most important in terms of species number and abundance. The maximum values of total microphytoplankton were observed at 0.5 m depth (46.2 × 10³ cells l⁻¹ at st. 3) in May as this was the month when the diatom Pseudo-nitzschia pungens bloomed. Chlorophyll (chl) a concentration ranged between 0.08 (August) and 0.78 μg l⁻¹ (February). May was another important month in which chlorophyll a increased (0.41–0.47 μg l⁻¹). Species diversity values (H′_log2) ranged from 1.66 bits (June, 20 m) to 4.11 bits (November, 0.5 m). The increase was attributed to a more balanced distribution of abundance among species. The amounts of nitrate + nitrite (0.6−3.7 μg-at N l⁻¹), phosphate (0.2−0.6 μg-at P l⁻¹) and silicate (0.7−2.5 μg-at Si l⁻¹) were recorded on each sampling occasion. Nutrient concentrations and chl a values of the research area were found to be poorer than those of the many other coastal areas in the northeastern Mediterranean. The mean atomic ratio of nitrogen to phosphorus varied from 1.3 (June) to 12.9 (February). This ratio was lower than the Redfield ratio of 16 for ocean phytoplankton, and phytoplankton was potentially limited by nitrogen for most of the months. The result of this study confirms and emphasizes the oligotrophic nature of the eastern Mediterranean.     

Fish species richness evaluation in Mexican coastal lagoons: a case study in the Gulf of Mexico

We analyze the origin of knowledge about fish species richness in the Tuxpan-Tampamachoco estuarine system, in Veracuz, México. A complete inventory of the fish species known to date for this system (N = 179) was elaborated from published lists and from sampling seagrass meadows of Tampamachoco Lagoon, which yielded 14 previously unknown species. When compared, the different lists showed a low similarity that may reflect differences in sampling methods and collecting strategies. Current data suggest that fish species richness in Mexican coastal lagoons (Gulf of Mexico) is not related with lagoon surface area, as has been suggested, but with the number of inventories available for each lagoon, being these a reflection of the sampling effort. A sampling design for the assessment of fish species richness in estuarine systems should consider: a) using the highest possible variety of sampling fishing gears, b) collecting in all microhabitat types and c) the preference of bimonthly or quarterly samplings for two or more years over monthly samplings in a single year.     

Variations in spatial and temporal distribution of Archaea in the North Sea in relation to environmental variables

The spatial and temporal distribution of pelagic Archaea was studied in the southern North Sea by rRNA hybridization, sequencing and quantification of 16S rRNA gene and membrane lipid analyses and related to physical, chemical and biological parameters to determine the factors influencing archaeal biogeography. A clear temporal variability was observed, with marine Crenarchaeota (Group I.1a) being relatively more abundant in winter and Euryarchaeota dominating the archaeal assemblage in spring and summer. Spatial differences in the lateral distribution of Crenarchaeota were also evident. In fact, their abundance was positively correlated with the copy number of the gene encoding the alpha subunit of crenarchaeotal ammonia monooxygenase (amoA) and with concentrations of ammonia, nitrate, nitrite and phosphorus. This suggests that most Crenarchaeota in the North Sea are nitrifiers and that their distribution is determined by nutrient concentrations. However, Crenarchaeota were not abundant when larger phytoplankton (>3 microm) dominated the algal population. It is hypothesized that together with nutrient concentration, phytoplankton biomass and community structure can predict crenarchaeotal abundance in the southern North Sea. Euryarchaeotal abundance was positively correlated with chlorophyll a concentrations, but not with phytoplankton community structure. Whether this is related to the potential of Euryarchaeota to perform aerobic anoxygenic phototrophy remains to be shown, but the conspicuous seasonal distribution pattern of Crenarchaeota and Euryarchaeota suggests that they occupy a different ecological niche.