Regime shifts in vegetation composition of Baltic Sea coastal lagoons

Vegetation communities, morphometric and water quality variables were sampled in 62 undisturbed coastal lagoons along a natural land-uplift gradient in the northern Baltic Sea. The lagoons had a morphological inlet threshold reflecting habitat isolation (i.e. the diminished connectedness of the lagoon to the sea) and located in different parts of the archipelago (corresponding to altering wave exposure). We used indirect multivariate methods to find changes in vegetation composition, and axes derived from the analyses were used in correlation and stepwise multiple regression analyses together with environmental variables. Habitat isolation proved to be the strongest predictor for vegetation composition. To identify significant change-points we used a method, which detects regime shifts by accounting for differences in chronological series and gives a regime shift index. Significant shifts in habitat isolation, average depth and total nitrogen were followed by shifts in species variables (i.e. dominant species, cover and species richness) and salinity, total phosphorus and turbidity. Of the species, especially Chara tomentosa showed a shift of high magnitude. Also a shift in wave exposure mirroring position of lagoons in the archipelago structured the species, but this division was seen only for less isolated lagoons. Our results suggest that lagoons are primarily structured by habitat isolation forming two structurally different regimes: one of less isolated lagoons dominated by a diverse array of vascular plants and marine algae of both outer and inner archipelago characters, and one with isolated lagoons dominated by high vegetation cover and dominance of Chara. In addition, vegetation seemed to stabilize water quality in the isolated regime by reducing turbidity and phosphorus, indicating a potential feed-back mechanism and a shift in structuring disturbances from abiotic to biotic factors with habitat isolation.     

Species diversity and distribution of aquatic macrophytes in the Northern Quark, Baltic Sea

The occurrence of aquatic macrophytes was studied in a northern transition area of the Baltic Sea; the Northern Quark, Gulf of Bothnia. In the area there is a gradual, marked change in environmental conditions, the most prominent of which is a decrease in salinity from 5.0% in the Bothnian Sea to 3.5% in the Bothnian Bay. In all, 40 species of macrophytes were observed; 10 fucophyceans, 10 bangiophyceans, 8 chlorophyceans, 3 charophyceans, 1 tribophycean, 1 nostocophycean, 6 phanerogams and 1 water moss. 26 of the observed species were of marine and 14 of lacustrine origin. There was a clear change in species composition and community structure from south to north over the area. The vegetation at the southernmost localities had a marine character, with belt-forming Fucus vesiculosus and a comparatively diverse flora of macroalgae. Further north, an ephemeral, lacustrine vegetation dominated by benthic diatoms and Cladophora aegagropila prevailed. The ratio marine: lacustrine species decreased from 4.2 to 1 when comparing a southern and a northern sub-area of the Northern Quark. The species observed include 57 % of the marine macrophytes noted in the Aland and Archipelago Seas (N Baltic Proper) during the past two decades. Two marine species, Aglaothamnion roseum and Ahnfeltia plicata , are reported for the first time from the Northern Quark. This comprises a northern extension of their distribution limit with approximately 300 km.     

Sulphide detoxification in Hediste diversicolor and Marenzelleria viridis, two dominant polychaete worms within the shallow coastal waters of the southern Baltic Sea

The polychaete worms Marenzelleria viridis (Verrill 1873) and Hediste diversicolor (O.F. Müller) form the main part of the macro-zoobenthos in soft-bottomed shallow inlets of the Baltic Sea. Due to high eutrophication within these waters the animals are exposed to low oxygen and high sulphide concentrations. Specimens of both species from a low salinity location (S 8 ‰) were compared concerning their physiological abilities in coping with this hostile environment. Sulphide detoxification occurred in both polychaetes even during severe hypoxia with the main end-product being thiosulphate. In absence of sulphide nearly no end-products of anaerobic metabolism were found in the worms during moderate hypoxia (pO₂=7 kPa). In presence of hydrogen sulphide, succinate, a sensitive indicator of anaerobic metabolism, was accumulated in higher amounts at low sulphide concentrations (0.3 mM) already. Oxygen consumption and ATP production was determined in isolated mitochondria of both species. Both polychaetes were able to perform enzymatic sulphide oxidation in the mitochondria at concentrations up to 50 μM. This process was coupled with oxidative phosphorylation. At least in M. viridis sulphide respiration was not completely inhibited by cyanide, suggesting an alternative oxidation pathway, which by-passes the cytochrome-c-oxidase. The two species did not differ in the rate of sulphide detoxification, but H. diversicolor produced about as twice as much ATP from mitochondrial sulphide oxidation. Differences in mitochondrial sulphide oxidation are probably related to the different life strategies of the worms.     

Epiphyte shading: Its role in resulting depth distribution of submerged aquatic macrophytes

The biomass of epiphytic algae increased more than the biomass of phytoplankton with increased nutrient availability (N, P, Si) in four lakes. Large epiphyte biomasses on the submerged macrophyte,Littorella uniflora, reduce light attenuation more than phytoplankton did. High values of shading by epiphytes correspond with a reduction in the maximum depth penetration ofLittorella by more than 1 meter.     

The palaeoenvironments of coastal lagoons in the southern Baltic Sea,I. The application of sedimentary Corg/N ratios as source indicators of organic matter

C/N ratios of organic matter sources in coastal lagoons and their drainage areas in the southern Baltic Sea region are investigated to determine the origin of organic matter (i.e. terrigenous or marine) in the sediments of the lagoons. These ratios are compared with δ¹³C values in the area. Terrestrial plants in the study area show C/N ratios between 10 and 85, whereas aquatic macrophytes have values in the range of 6 to 44. Peat and shore vegetation have C/N ratios between 15 and 16, and plankton shows values close to 6. In conjunction with δ¹³C values, the C/N ratios in the sediments allow the reconstruction of the Holocene palaeoenvironments and palaeoecology in the lagoons (Oder Estuary and Greifswalder Bodden). Distinct stages in the development of the water bodies, resulting from sea level changes in the region, can be derived: post-glacial lake stages with sandy sedimentation, lacustrine phases with high autochthonous productivity, terrestrial stages with peat formation, sedimentation as a result of marine transgression, and brackish sedimentation after the formation of sand spits and barrier islands. The application of C_org/N_tot ratios instead of C_org/N_org does not have any significant influence on the interpretation of the data and may thus be used in the coastal sediments of the southern Baltic Sea after having applied the test procedures presented in this study.     

Climate-related differences in the dominance of submerged macrophytes in shallow lakes

It has been suggested that shallow lakes in warm climates have a higher probability of being turbid, rather than macrophyte dominated, compared with lakes in cooler climates, but little field evidence exists to evaluate this hypothesis. We analyzed data from 782 lake years in different climate zones in North America, South America, and Europe. We tested if systematic differences exist in the relationship between the abundance of submerged macrophytes and environmental factors such as lake depth and nutrient levels. In the pooled dataset the proportion of lakes with substantial submerged macrophyte coverage (> 30% of the lake area) decreased in a sigmoidal way with increasing total phosphorus (TP) concentration, falling most steeply between 0.05 and 0.2 mg L⁻¹. Substantial submerged macrophyte coverage was also rare in lakes with total nitrogen (TN) concentrations above 1–2 mg L⁻¹, except for lakes with very low TP concentrations where macrophytes remain abundant until higher TN concentrations. The deviance reduction of logistic regression models predicting macrophyte coverage from nutrients and water depth was generally low, and notably lowest in tropical and subtropical regions (Brazil, Uruguay, and Florida), suggesting that macrophyte coverage was strongly influenced by other factors. The maximum TP concentration allowing substantial submerged macrophyte coverage was clearly higher in cold regions with more frost days. This is in agreement with other studies which found a large influence of ice cover duration on shallow lakes' ecology through partial fish kills that may improve light conditions for submerged macrophytes by cascading effects on periphyton and phytoplankton. Our findings suggest that, in regions where climatic warming is projected to lead to fewer frost days, macrophyte cover will decrease unless the nutrient levels are lowered.     

The chemical character and burial of phosphorus in shallow coastal sediments in the northeastern Baltic Sea

The chemical composition and vertical distribution of sediment phosphorus (P) in shallow coastal sediments of the northeastern Baltic Sea (BS) were characterized by sequential extraction. Different P forms were related to chemical and physical properties of the sediments and the chemistry of pore water and near-bottom water. Sediment P composition varied among the sampling sites located in the Archipelago Sea (AS) and along the northern coast of the Gulf of Finland (GoF): the organic rich sites were high in organic P (OP), while apatite-P dominated in the area affected by sediment transportation. Although the near-bottom water was oxic, the sediments released P. Release of P was most pronounced at the site with high sediment OP and reduced conditions in the sediment-water interface, indicating that P had its origins in organic sources as well as in reducible iron (Fe) oxyhydroxides. The results suggest that even though these coastal areas are shallow enough to lack salinity stratification typical for the brackish BS, they are vulnerable to seasonal oxygen (O₂) depletion and P release because of their patchy bottom topography, which restricts mixing of water. Furthermore, coastal basins accumulate organic matter (OM) and OP, degradation of which further diminishes O₂ and creates the potential for P release from the sediment. In these conditions, an abundance of labile OP may cause marked efflux of P from sediment reserves in the long-term.     

Temperature and light: The determining factors in maximum depth distribution of aquatic macrophytes in Ontario,Canada

In stratified lakes with high light penetration, the maximum depth at which macrophytes occur is frequently limited by temperature. At this depth a variety of species may be found. On the other hand, when the clarity of water limits the light penetration and the temperature at depth is sufficient for good plant growth, the plants occurring at greatest depth are ones that do not require photosynthetic oxygen for root growth. Such plants include the Charales, Isoetes, Utricularia and Ceratophyllum.     

Effects of subsidized spiders on coastal food webs in the Baltic Sea area

The aim of this study was to examine top-down effects of cursorial spiders in subsidized coastal food webs. Top-down effects were examined by selectively removing cursorial spiders, mainly wolf spiders, from small islands (26–1834 m²) during 2004–2007. The removal success varied among islands and years, and spider densities were reduced by 30–65%. To examine treatment effects, arthropods were sampled using a vacuum sampling device at three occasions each summer. The densities of other arthropod predators, especially web spiders and carabids, were higher on islands where cursorial spiders had been removed compared to control islands. This treatment effect probably occurred through a combination of competitive release and reduced intraguild predation from cursorial spiders. No treatment effects were found on herbivore or detritivore densities and plant biomass. This lack of effect may either be because spiders indeed have fairly weak effects on herbivore and detritivore densities on Baltic shorelines or that the removal success of spiders was insufficient for observing such effects. Treatment effects may also be weak because negative effects exerted by spiders on herbivore and detritivore populations were balanced by increased predation by insect predators.     

Ecological classification of macroalgae and angiosperm communities of inner coastal waters in the southern Baltic Sea

A classification system of macroalgae and angiosperms for the inner coastal waters of the Baltic coast of Mecklenburg-Vorpommern (Germany) was developed according to the guidelines of the European Water Framework Directive. These guidelines ask for a five-step classification scheme (high, good, moderate, poor, bad) of the ecological state based on submerged macrophytes.The approach is based on the comparison of the occurrence and distribution of plant communities in 1999–2003 with historical data of submerged macrophytes and their community structure. Due to the lack of appropriate data, historical depth distribution limits of plant communities were calculated based on modelled historical underwater light climate.Despite the fact of very heterogeneous salinities (0.5–20.0 practical salinity units, psu) and substrate types (silty–sandy) in the study area, two general main criteria were identified as appropriate indicators of eutrophic degradation of the inner coastal waters: (1) the lower distribution depth of plant communities and (2) the loss of charophyte-dominated plant communities. Based on these two criteria, a five-step classification scheme is proposed for the characterisation of the ecological state. The depth limit boundaries between the ecological classes were calculated by 5, 25, 50 and 75% reduction of the water transparency. Based on this calculation, the ecological status is described by the gradual upward shift of the vegetation depth limit. Complementary to the decrease of depth limits the loss of charophyte-dominated plant communities characterises the border between moderate and poor ecological conditions.     

Spatio-temporal distribution and production of calanoid copepods in the central Baltic Sea

The aim of our study was the exploration of species-specificdistribution and production patterns of dominant copepods inthe Central Baltic Sea (Bornholm Basin). Spatio-temporal distribution,egg and secondary production were studied by means of net-samplingand egg production experiments from April to August 1999. Verticaland horizontal distribution patterns appeared to be linked witheach other and were primarily species-dependent. Both Acartiaspp. and Temora longicornis preferentially inhabited the upper30 m of the water column and the shallower marginal regionsof the Bornholm Basin. In contrast, the C4–5 and C6 ofPseudocalanus spp. preferentially inhabited the halocline region(50–70 m) and the deep central part of the area. Observeddifferences in horizontal distribution among these three copepodsappear to result from different depth preferences and depth-dependentwater circulation patterns. Egg production rates (EPR) clearlyexhibited seasonal differences with maximum EPR in May and minimumEPR in July. Mean EPR was significantly different between species,being highest in Acartia spp. and lowest in Pseudocalanus spp.The variability in EPR was related to differences in hydrographyand modelled food environment.     

Bacterial interactions in the rhizosphere of seagrass communities in shallow coastal lagoons

Rooted phanerogam communities in the shallow intertidal and subtidal coastal zone represent productive and healthy ecosystems. Inorganic nutrients are assimilated into seagrass biomass. Much of the organic matter resulting from moribund seagrass is rapidly mineralized, principally by bacteria. The microbial community of the rhizosphere is also highly active due to the supply of organic matter released during photosynthesis. This active sediment community plays an important role through carbon, nitrogen and phosphorous cycling in maintaining the stability and productivity of seagrass meadows. Over the last two decades, however, seagrass meadows in European coastal areas have declined due to increasing pollution. As eutrophication advances a trasition occurs from rooted phanerogram dominated communities to planktonic algal blooms and/or cyanobacterial blooms. Such changes represent the decline of a stable, high biodiversity habitat to an unstable one dominated by a few species. These changes of community structure can occur rapidly once the internal nutrient and organic matter control cycles are exceeded. A field investigation was undertaken to establish the spatial distribution of bacterial populations of Zostera noltii colonized and uncolonized sediment in the Bassin d'Arcachon, France. Bacteria were enumerated using both plate count and MPN techniques for different functional groups as well as determining the total bacterial populations present. Nitrogen fixation, ammonification, sulphate reduction rates, as well as alkaline phosphatase activity were also determined. Colonization of the Z. noltii roots and rhizomes was studied by light and scanning electron microscopy. Results confirmed that higher bacterial populations were present in the rhizosphere of Z. noltii compared to uncolonized sediments. Furthermore, electron microscopy identified the rhizome as the main site of colonization for a diverse range of morphological groups of bacteria. Sulphate reducing bacteria were identified as the key group of bacteria involved in N-fixation in the rhizosphere of Z. noltii. The data will be discussed in relation to the role played by the rhizosphere microflora in supplying and mobilising nutrients in Z. noltii.     

Long- and short-term dynamics of submerged macrophytes in two shallow eutrophk lakes

1. Periods with clear water and abundant submerged vegetation have alternated with periods of turbid water and sparse vegetation during recent decades in Lake Tåkern and Lake Krankesjön, two shallow, calcium-rich, moderately eutrophic lakes in southern Sweden, Between 1983 and 1991, submerged vegetation (predominant species: Chara tomentosa, Nitellopsis obtusa, Myriophyllum spicatum) covered about 50% of the open lake area in Lake Tåkern. In Lake Krankesjön, submerged vegetation was sparse during 1983–84, but increased continuously in the following years and covered about 50% of the open lake area by 1990 and 1991. Potamogeton pectinatus was the first species to expand in Lake Krankesjön, but was later replaced by C. tomentosa. 2. During 1983–84, turbidity was high in Lake Krankesjön, which indicated that submerged macrophytes were light-limited. During 1986–91, there was a negative correlation between the areal coverage of charophytes and angiosperms, indicating that competition for space had become an important limiting factor. The same negative correlation was found in Lake Tåkern for 1983–91. 3. Charophytes had much higher biomass per unit area than angiosperms in both lakes and reduced water movement considerably. This was probably one reason for the increase of water transparency in Lake Krankesjön during the spatial expansion of these plants. Charophytes also stored large amounts of phosphorus and nitrogen, Charophytes are probably superior competitors for both space and nutrients and thus have competitive advantage over angiosperms in this lake type. 4. In Lake Krankesjön, both P. pectinatus and C. tomentosa were negatively affected by high water level during the growing period. Total disappearance of submerged vegetation occurred in both lakes after catastrophic events (dry-out during summer or mechanical damage by ice) caused by extremely low water level. Changes in water level are thus one of the most important reasons for among-year fluctuations in areal coverage of submerged macrophytes in these lakes.     

Vertical distribution and chemical character of sediment phosphorus in two shallow estuaries in the Baltic Sea

The vertical distribution of various phosphorus (P) forms and their relation to physico-chemical properties of estuary sediment material were studied to better understand the potential release and burial of P. Core samples were taken from two dissimilar estuaries in the Baltic Sea: one in the Archipelago Sea (AS) and one in the Gulf of Finland (GoF). The P reserves were characterized by a sequential extraction procedure including the analysis of simultaneously dissolved elements in two extraction steps. The sediment material was also analysed for particle size distribution and total elements. In addition, several environmental variables were determined. The occurrence of the various forms of P varied with sediment depth among different sites. Reductant soluble, iron (Fe) bound P was the most dynamically changing P form in the sediment, while P bound to other metal oxides and apatite-P were the most stable fractions. High sedimentation rate was a dominating factor for sediment P burial. In addition, the content of organic matter, the amount of erosion-transported sorption components, and the oxygen (O₂) conditions in the near-bottom water were important determinants of the behaviour of sediment P. The results indicate that, over the long term, both estuaries have acted as sinks for deposited P and restricted the transport of P to the AS and the open GoF, thereby partly alleviating the eutrophication process.     

Selective predation by herring and mysids, and zooplankton community structure in a Baltic Sea coastal area

In lakes, fish and invertebrate predation are recognized asstrong structuring forces on zooplankton communities. The objectof this investigation was to study whether predation has a similarpotential in a coastal area of the brackish Baltic Sea and ifit could explain observed differences in zooplankton communitystructure between a reference area and an eutrophied area. Speciescomposition and daily vertical migration of zooplankton andzooplanktivores, and the diets of the latter, were studied inJuly and August at two 30 m deep stations differing in primaryproductivity. The biomass of zooplankton >35 µm wasdominated by copepods, but cladocerans and rotifers also occurredin significant numbers. The dominating zooplanktivores wereherring (Clupea harengus) and the mysid shrimp Mysis mixta.They fed almost exclusively on zooplankton, mainly copepods,and their estimated food consumption equalled or exceeded thesummer copepodite production. The structure of the zooplanktoncommunity cannot be explained by effects of predation or byfeeding conditions alone. Increases in rotifer and cladoceranabundances with increased primary production suggests effectsof food supply. However, a generally rapid decline in the annualsummer peak of cladocerans may be caused by predation. The totalabundance of copepods did not increase with improved feedingconditions, but there was a shift in species dominance. Thecopepod Ewytemora affinis hirundoides, which was intensivelypreyed upon, increased with increased phytoplankton production,while Acartia bifilosa and/or A.longiremis, which was less eaten,decreased. Predation may explain a pronounced daily verticalmigration of the most predated copepods. They occurred in deeperwater during the day, when the visually feeding herring wereactive, and moved closer to the surface at night when M.mixtaleft the bottom, to forage in the water column.     

Abundance, depth distribution, and composition of aerobic bacteriochlorophyll a-producing bacteria in four basins of the central Baltic Sea

The abundance, vertical distribution, and diversity of aerobic anoxygenic phototrophic bacteria (AAP) were studied at four basins of the Baltic Sea. AAP were enumerated by infrared epifluorescence microscopy, and their diversity was analyzed by using pufM gene clone libraries. In addition, numbers of CFU containing the pufM gene were determined, and representative strains were isolated. Both approaches indicated that AAP reached maximal abundance in the euphotic zone. Maximal AAP abundance was 2.5 x 10(5) cells ml(-1) (11% of total prokaryotes) or 1.0 x 10(3) CFU ml(-1) (9 to 10% of total CFU). Environmental pufM clone sequences were grouped into 11 operational taxonomic units phylogenetically related to cultivated members of the Alpha-, Beta-, and Gammaproteobacteria. In spite of varying pufM compositions, five clones were present in all libraries. Of these, Jannaschia-related clones were always found in relative abundances representing 25 to 30% of the total AAP clones. The abundances of the other clones varied. Clones potentially affiliated with typical freshwater Betaproteobacteria sequences were present at three Baltic Sea stations, whereas clones grouping with Loktanella represented 40% of the total cell numbers in the Gotland Basin. For three alphaproteobacterial clones, probable pufM phylogenetic relationships were supported by 16S rRNA gene analyses of Baltic AAP isolates, which showed nearly identical pufM sequences. Our data indicate that the studied AAP assemblages represented a mixture of marine and freshwater taxa, thus characterizing the Baltic Sea as a "melting pot" of abundant, polyphyletic aerobic photoheterotrophic bacteria.