Pauli Snoeijs

Long-term changes in the macroalgal vegetation of the inner Gullmar Fjord, Swedish Skagerrak coast

We examined long‐term changes in the macroalgal vegetation at Stora Bornö Island in the inner Gullmar Fjord on the Swedish Skagerrak coast. This was made possible by access to a 1941 diving investigation. The same sites were reinvestigated in 1998. Community composition and depth distributions of species were compared and changes were analyzed with focus on functional groups (size, thallus shape, and life‐history traits). We discovered a significant decrease in the depth extension of macroalgal species and a dramatic decline of species richness in the lower littoral (below 16 m of depth) compared with 57 years earlier. Ordination analysis revealed that there was a significant difference in the community composition between the two study periods. In general, small (<10 cm), thin, filamentous, and aseasonal ephemerals increased in relative abundance, whereas larger (>10 cm), coarsely branched, and perennial algae decreased. Calibrations of individual species to local sediment cover, using canonical correspondence analysis, indicated that part of the change in species composition was related to sediment load. Furthermore, large‐scale climate differences (NAO Winter Index) between the study periods indicated a higher impact of Baltic Sea and Kattegat water in the nutrient dynamics of the fjord in the 1998 study. We concluded that the observed long‐term changes in the macroalgal community at Stora Bornö Island were consistent with an increased nutrient availability.

Long-term changes in the sublittoral zonation of brown algae in the southern Bothnian Sea

The algal vegetation of five diving profiles situated at exposed sites of the outer Oregrund archipelago, southern Bothnian Sea, Sweden, was investigated by SCUBA diving in 1996. The vegetation of the same profiles had been described earlier in 1943-4 and 1984. In the present study, special attention was paid to the occurrence of the structurally important belt-forming brown algae Fucus vesiculosus and Sphacelaria arctica in comparison with the situation 52-53 and 12 years earlier. Prominent differences between the 1940s and 1996 were found. The weighted average depth of the F. vesiculosus belt in 1996 was about 1.7 m shallower than in the 1940s, and the lower distribution limit of this species was about 2.5 m shallower. At its lower end the Fucus belt was replaced by epilithic red algae, mainly Furcellaria lumbricalis, Rhodomela confervoides and Polysiphonia fucoides. These results were similar to those of the 1984 study and show neither an improvement nor a deterioration of the belt during the past 12 years. In four of the five diving profiles, a conspicuous belt of S. arctica occurred in the lower sublittoral, similar to that observed in the 1940s, but in one of the five profiles the S. arctica belt was heavily impoverished. These results contrast with the 1984 study which reported that S. arctica had practically disappeared from all profiles. The epiphytic cover of annual filamentous algae, dominated by Pilayella littoralis and Ceramium gobii, was higher in 1996 than in 1943-4. The results of this comparative study are discussed in relation to the general eutrophication of the Baltic Sea.

THE IMPORTANCE OF DIATOM CELL SIZE IN COMMUNITY ANALYSIS1

The large variation in size and shape in diatoms is shown by morphometric measurements of 515 benthic and pelagic diatom species from the Baltic Sea area. The largest mean cell dimension (mostly the apical axis) varied between 4.2 and 653 μm, cell surface area between 55 and 344,000 μm2, and cell volume between 21 and 14.2 × 106μm3. The shape‐related index, length to width ratio, was between 1.0 and 63.3 and the shape‐ and size‐related index, surface area to volume ratio, was between 0.02 and 3.13. Diatom community analysis by multivariate statistics is usually based on counts of a fixed number of diatom valves with species scores irrespective of cell size. This procedure underestimates the large species for two reasons. First, the importance of a species with higher cell volume is usually larger in a community. Second, larger species usually have lower abundances and their occurrence in the diatom counts is stochastic. This article shows that co‐occurring small and large diatom species can respond very differently to environmental constraints. Large epiphytic diatoms responded most to macroalgal host species and small epiphytic diatoms most to environmental conditions at the sampling site. Large epilithic diatoms responded strongly to salinity, whereas small epilithic diatoms did so less clearly. The conclusion is that different scale‐dependent responses are possible within one data set. The results from the test data also show that important ecological information from diatom data can be missed when the large species are neglected or underestimated.

Effects of temperature on the production of hydrogen peroxide and volatile halocarbons by brackish-water algae

Marine algae produce volatile halocarbons, which have an ozone-depleting potential. The formation of these compounds is thought to be related to oxidative stress, involving H2O2 and algal peroxidases. In our study we found strong correlations between the releases of H2O2 and brominated and some iodinated compounds to the seawater medium, but no such correlation was found for CHCl3, suggesting the involvement of other formation mechanisms as well. Little is known about the effects of environmental factors on the production of volatile halocarbons by algae and in the present study we focused on the influence of temperature. Algae were sampled in an area of the brackish Baltic Sea that receives thermal discharge, allowing us to collect specimens of the same species that were adapted to different field temperature regimes. We exposed six algal species (the diatom Pleurosira laevis, the brown alga Fucus vesiculosus and four filamentous green algae, Cladophora glomerata, Enteromorpha ahlneriana, E. flexuosa and E. intestinalis) to temperature changes of 0-11 degrees C under high irradiation to invoke oxidative stress. The production rates, as well as the quantitative composition of 16 volatile halocarbons, were strongly species-dependent and different types of responses to temperature were recorded. However, no response patterns to temperature change were found that were consistent for all species or for all halocarbons. We conclude that the production of certain halocarbons may increase with temperature in certain algal species, but that the amount and composition of the volatile halocarbons released by algal communities are probably more affected by temperature-associated species shifts. These results may have implications for climatic change scenarios.

DISTRIBUTION OF EPIPHYTIC DIATOM SPECIES COMPOSITION, DIVERSITY AND BIOMASS ON DIFFERENT MACROALGAL HOSTS ALONG SEASONAL AND SALINITY GRADIENTS IN THE BALTIC SEA

In spring 1990 and 1991, some weeks after ice-break, macroalgae and their epiphytes were sampled from 8 areas along the stable salinity gradient of the Baltic Sea from the southern Baltic Sea proper (8‰) to the northern Bothnian Sea (5‰). The macroalgal hosts included the brown alga Pilayella littoralis (L.) Kjellm., the red alga Ceramium gobii Waern, and the green alga Cladophora glomerata (L.) Kutz. In addition, seasonal succession was studied from monthly samples throughout one growing season (March-November 1991) at one of the sampling sites (5‰ salinity). Host preference was highest in spring when the diatoms were newly colonizing the rapidly growing macroalgae, and higher in the Baltic Sea proper (higher salinity) than in the Bothnian Sea (lower salinity). Biomass was generally highest on Pilayella and lowest on Cladophora, whereas community diversity was generally highest on Ceramiutn and lowest on Pilayella. The differences in epiphytic diatom community composition and biomass between the three mac...

Patterns of macroalgal diversity, community composition and long-term changes along the Swedish west coast

This study analyses the complicated patterns of vertical distribution of the macroalgal vegetation in an area where brackish and marine waters meet and mix. Variables used to record vegetation characteristics are algal cover, species composition and diversity. The data set includes 64 diving profiles, all from sites exposed to wave action, along a ca. 260 km long coastline. The profiles belong to four categories: coastal sites in the Skagerrak (more marine), coastal sites in the Kattegat (more brackish), coastal sites in the Kattegat after a toxic phytoplankton bloom, and submerged offshore stone reefs in the Kattegat. The highest species diversity was found at the reefs, which are not affected directly by land runoff. At the reefs the 18 most common perennial species penetrate 2–11 m (on average 5.5 m) deeper than at the coastal sites. The virtual absence of sedimentation, and thus the availability of substratum, at the reefs may explain the differences so that the lower limit for the algae is determined by light penetration or by recruitment problems caused by strong currents at the reefs, whereas sedimentation limits the settlement of algae in coastal sites. Ordination analysis based on species composition reveals that the major environmental gradients structuring the algal vegetation in the Kattegat and the Skagerrak are salinity and water depth. The large data set of this study made it possible to quantify the downward dislocation of Atlantic intertidal species to the sublittoral along the Swedish west coast. For example, the mean upper limit of Corallina officinalis is 2 m in the Skagerrak but 12.5 m in the Kattegat and the mean occurrence interval of Fucus serratus is 0.9–2.7 m in the Skagerrak, but 1.1–6.3 m in the Kattegat. This downward dislocation is suggested to be the result of decreased competition when species successively disappear with lower salinity. Comparisons of the present studys results with those of previous investigations show that eight common red algal species have moved upwards compared to the situation before the large-scale eutrophication started in the 1960s, e.g. Cystoclonium purpureum and Polysiphonia elongata by ca. 8 m, Phycodrys rubens and Delesseria sanguinea by ca. 5 m. A toxic phytoplankton bloom affected macroalgal community composition on the whole only slightly, but it had a negative effect on algal cover and species richness below a water depth of ca. 5 m, the algae were visibly damaged and the lower vegetation limit temporarily moved upwards.

THE NATURAL LIFE CYCLE IN WILD POPULATIONS OF DIATOMA MONILIFORMIS (BACILLARIOPHYCEAE) AND ITS DISRUPTION IN AN ABERRANT ENVIRONMENT1

We studied how size variation in populations of Diatoma moniliformis Kütz. was influenced by environmental effects on the diatom life cycle. One of the two populations sampled monthly in the northern Baltic Sea grew under natural conditions; the other population was in a cooling water discharge channel of a nuclear power plant, where the temperature and flow rate of the water were artificially higher. The life cycle was synchronous at the natural site, with sexual reproduction occurring in the winter; most of the initial cells were found in March‐April. After this, a reduction in cell size occurred, and the vegetative life cycle consisted of two parts. During the first part, cell volume decreased, whereas the surface area to volume ratio increased, and during the second part of the cycle, both of these parameters decreased. No direct evidence was found for the existence of a supra‐annual life cycle in D. moniliformis, as convincing modes fm large cells were lacking in the size‐frequency distributions. It was concluded from extrapolations of the data that the natural life cycle of D. moniliformis probably lasts 2 or 3 years. The changes in cell proportions during the life cycle fit well with annual growth cycles of D. moniliformis at the natural site (i.e. the cells had high surface area to volume ratios during the period of optimal growth in late spring [May‐June]). At the site affected by cooling water discharge, the synchronization of the natural life cycle was disrupted, but some seasonal size variation did occur. Under natural conditions, auxosporulation is probably triggered by a combination of small cell size, low water temperature (0–3° C), and rapidly increasing light intensity or daylength in late winter to early spring. When these conditions were not met (e.g. at the heated site, the required low temperature was absent), auxosporulation did not occur simultaneously. This paper also presents scanning electron photomicrographs showing the typical shape and fine structure of the initial cell of D. moniliformis. These cells are semispherical in cross section, possess a pronounced curvature along the longitudinal axis, and are bent in the perualvar plane.

UPTAKE OF INORGANIC CARBON BY CLADOPHORA GLOMERATA (CHLOROPHYTA) FROM THE BALTIC SEA1

Carbon uptake in the green macroalga Cladophora glomerata (L.) Kütz. from the brackish Baltic Sea was studied by recording changes in pH, alkalinity, and inorganic carbon concentration of the seawater medium during photosynthesis. The use of specific inhibitors identified three uptake mechanisms: 1) dehydration of HCO3− into CO2 by periplasmic carbonic anhydrase, followed by diffusion of CO2 into the cell; 2) direct uptake of HCO3− via a 4,4′‐diisothiocyanato‐stilbene‐2,2′‐disulfonate‐sensitive mechanism; and 3) uptake of inorganic carbon by the involvement of a vanadate‐sensitive P‐type H+‐ATPase (proton pump). A decrease in the alkalinity of the seawater medium during carbon uptake, except when treated with vanadate, indicated a net uptake of the ionic species contributing to alkalinity (i.e. HCO3−, CO32−, and OH−) from the medium, where OH− influx is equivalent to H+ efflux. This would suggest that the proton pump is involved in HCO3− transport. We also show that the proton pump can be induced by carbon limitation. The inducibility of carbon uptake in C. glomerata may partly explain why this species is so successful in the upper littoral zone of the Baltic Sea. Usually, carbon limitation is not a problem in the upper littoral of the sea. However, it may occur frequently within dense Cladophora belts with high photosynthetic rates that create high pH and low carbon concentrations in the algas microenvironment.

Long-term changes of macroalgal vegetation in the Skagerrak area

The algal vegetation at three rocky-shore localities on the Swedish Skagerrak coast with different environmental conditions was studied in 1960–1961 by SCUBA diving. The same localities were revisited in the summer of 1997, using the same methods for recording the vegetation. Detailed descriptions of the vegetation profiles are presented to allow the recording of future changes in these profiles, and extensive ecological herbaria are kept of both the 1960–1961 and the 1997 investigations. The abundance and depth distribution of 78 macroalgal species were recorded in 1997 and community composition was compared with that of the early 1960s. Increases of perennial red algae with delicate foliaceous thalli (Delesseria sanguinea, Phycodrys rubens) were conspicuous at the two localities most exposed to wave action, whereas perennial red algae with tougher foliaceous thalli (Phyllophora truncata and Phyllophora pseudoceranoides) prevailed at the more sheltered locality with most sedimentation. It is hypothesised that increased abundances of delicate species with a large growth potential are caused by eutrophication, but that this effect may be counteracted when eutrophication results in a high load of sedimentation. Tougher species are designed better to withstand a heavy load of sediment. Other perennial red algae with tougher thalli (Chondrus crispus, Furcellaria lumbricalis) also decreased at the exposed sites, but not at the site with most sedimentation, possibly because of lack of competition from D. sanguinea and P. rubens. The abundance of filamentous algae had increased since the early 1960s at all three localities, which may be attributed to eutrophication as well, but no changes were observed in the large perennial brown algae. Decaying loose-lying algae occurred at all localities, but algal mat formation was only abundant at the most sheltered locality. The observed changes in the attached algal vegetation of the exposed sites may be an indication of a general large-scale eutrophication of the Skagerrak.

Density-dependent patterns of thiamine and pigment production in the diatom Nitzschia microcephala

In the present study we investigate how intraspecific (density-dependent) competition for nutrients by the diatom Nitzschia microcephala affects the level of oxidative stress in the algal cells as well as their production of pigments and thiamine. N. microcephala was grown in three different densities until the stationary growth phase was reached. Throughout the experiment, growth rate was negatively related to cell density. Superoxide dismutase activity, protein thiol, and diatoxanthin concentrations indicated increasing oxidative stress with increasing cell density, which was most probably caused by nutrient depletion of the medium. Pigment contents per cell (except for diatoxanthin) decreased with increasing cell density. N. microcephala was able to synthesize thiamine and its thiamine content per cell increased in concert with cell density. In comparison, the dinoflagellate Amphidinium carterae was unable to synthesize thiamine. These results suggest that cells of N. microcephala subjected to higher competition and lower growth rates have a lower carotenoid content and a higher thiamine content. If such responses would occur in nature as well, eutrophication (higher cell densities) may alter the quality of microalgae as food items for higher trophic levels not only by species shifts in the phytoplankton, but also by changes in the cellular nutritional value within species.