Jonne Kotta

Hypoxia Is Increasing in the Coastal Zone of the Baltic Sea

Hypoxia is a well-described phenomenon in the offshore waters of the Baltic Sea with both the spatial extent and intensity of hypoxia known to have increased due to anthropogenic eutrophication, however, an unknown amount of hypoxia is present in the coastal zone. Here we report on the widespread unprecedented occurrence of hypoxia across the coastal zone of the Baltic Sea. We have identified 115 sites that have experienced hypoxia during the period 1955–2009 increasing the global total to ca. 500 sites, with the Baltic Sea coastal zone containing over 20% of all known sites worldwide. Most sites experienced episodic hypoxia, which is a precursor to development of seasonal hypoxia. The Baltic Sea coastal zone displays an alarming trend with hypoxia steadily increasing with time since the 1950s effecting nutrient biogeochemical processes, ecosystem services, and coastal habitat.

Ecological consequences of biological invasions: three invertebrate case studies in the north-eastern Baltic Sea

Population dynamics and ecological impacts of the cirriped Balanus improvisus, the polychaete Marenzelleria neglecta and the cladoceran Cercopagis pengoi were investigated in the north-eastern Baltic Sea. After an increase during the first decade of invasion, the density of M. neglecta and C. pengoi declined afterwards. The studied abiotic environmental variables did not explain the interannual variability in the seasonal cycles of M. neglecta and C. pengoi indicating that the species are at their initial phase of invasion. The population dynamics of B. improvisus was best described by water temperature. B. improvisus promoted the growth of the green alga Enteromorpha intestinalis. M. neglecta enhanced the content of sediment chlorophyll a and reduced growth and survival of the polychaete Hediste diversicolor and growth of the amphipod Monoporeia affinis. Concurrent with the invasion of C. pengoi the abundance of small-sized cladocerans declined, especially above the thermocline. C. pengoi had become an important food for nine-spined stickleback, bleak, herring and smelt.

A meta-analysis of seaweed impacts on seagrasses: Generalities and knowledge gaps

Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than ‘rooted’ seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that ‘rooted’ seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.

Major Changes in Macroalgae Community Composition Affectthe Food and Habitat Preference of Idotea baltica

Followed by the massive increase in the biomass of Pilayella littoralis, Fucus vesiculosus has almost disappeared in Koiguste Bay, north-eastern Baltic Sea during 1995‐1998. Consequently, Idotea baltica, which previously fed on Fucus, switched to new food items ‐ Furcellaria lumbricalis and Pilayella littoralis. Idotea grazed more Pilayella than Furcellaria whereas Furcellaria was highly preferred as a habitat. Maximum 2.2% and 4.7% of the production of Furcellaria and Pilayella were removed by grazing. We conclude that the grazing pressure by Idotea is not sufficient to control the outbreak of Pilayella. However, Idotea has an advantageous effect on the persistence of Furcellaria by preferentially removing fast growing epiphyte Pilayella.

‘In vitro’ and ‘in situ’ decomposition of nuisance macroalgae Cladophora glomerata and Pilayella littoralis

The decomposition of two macroalgal species Cladophora glomerata (CHLOROPHYTA) and Pilayella littoralis (PHAEOPHYTA) was studied in the laboratory and field conditions. These species are known to cause the extensive ‘macroalgal blooms’ in the whole coastal range of the Baltic Sea. The objective of the experiments was to determine decomposition rates of the macroalgae, follow the changes in tissue nutrient content and validate the role of benthic invertebrates in this process. In the laboratory conditions, the differences in the decomposition rates of the algae were mainly due to the oxygen conditions. The weight loss of C. glomerata was slightly higher in anaerobic conditions than in aerobic conditions. If 99% of initial dry weight of P. littoralis was lost in aerobic conditions then only 20% was lost in anaerobic conditions. In general, the loss of phosphorus and nitrogen in algal tissues followed the weight loss. As an exception, the amount of nitrogen changed very little during the decomposition of C. glomerata. In field conditions, the photosynthetic activity exceeded the decomposition rate of C. glomerata at lower temperatures in spring. The decomposition of P. littoralis was estimated at 49% of its initial dry weight. The addition of benthic invertebrates had no effect on the decomposition process. In summer, the decomposition rates were estimated at 65% for C. glomerata and 68% for P. littoralis being in the same order of magnitude as observed in laboratory conditions. If the decomposition of C. glomerata was faster at the end of the experiment, the most significant losses of weight of P. littoralis took place during the first 2 weeks of deployment. Idotea baltica significantly contributed to the loss of C. glomerata. The decomposition rate of P. littoralis was reduced by the presence of Mytilus edulis and increased by Gammarus oceanicus.

Competition for food between the introduced polychaete Marenzelleria viridis (Verrill) and the native amphipod Monoporeia affinis Lindstrom in the Baltic Sea

Since 1985 the spionid polychaete Marenzelleria viridis (Verrill) has invaded large parts of the Baltic Sea. In deeper soft-bottom habitats (>10 m) a marked long-term decrease of the native amphipod Monoporeia affinis has been noted and is presently associated with the establishment of the polychaete. One plausible explanation is that the polychaetes and the amphipods are competing for food resources as both are deposit feeding animals and likely to share similar food resources. Interspecific competition for food between the polychaete and the amphipod was studied in a laboratory experiment. Two year classes (0y+,1y+) of the amphipods were kept at various densities, with and without added food resources, with and without the polychaete, in microcosms with sediment and continuous supply of cooled water for 2 months. The polychaetes did not have any effect on mortality in the amphipods. 4-way ANOVA showed that food addition, density of amphipods and presence of the polychaete had a significant effect on the growth of amphipods of different age classes. 1y+ amphipods showed increased growth with added food and this increase was density-dependent in the absence of the polychaetes but not in their presence. The polychaetes reduced the growth of 1y+ amphipods at natural densities (2000 ind m−2) by 60%, but had no clear effects on the growth of juveniles. It is concluded that lower amphipod growth in the presence of M. viridis was caused by competition for food and is likely to affect the population of M. affinis in deep soft-bottom habitats of the northern Baltic Sea.

Seasonal changes in situ grazing of the mesoherbivores Idotea baltica and Gammarus oceanicus on the brown algae Fucus vesiculosus and Pylaiella littoralis in the central Gulf of Finland, Baltic Sea

The in situ grazing experiments were performed in the shallow water rocky habitat of the northern Baltic Sea during ice-free season 2002. In the experiments the effects of algal species and choice on the grazing of the mesoherbivores Idotea baltica (Pallas) and Gammarus oceanicus Segerstrale were tested. Salinity, temperature, concentration of nutrients in water and macroalgae and net production of macroalgae were considered as random effects in the analysis. The invertebrate feeding rate was mainly a function of the net photosynthetic activity of Pylaiella littoralis (L.) Kjellman and Fucus vesiculosus L. Feeding rate increased significantly with decreasing algal photosynthetic activity. When the two algal species were incubated together invertebrates fed primarily on P. littoralis. Low selectivity towards P. littoralis coincided with its high photosynthetic activity. The presence of F. vesiculosus did not modify the invertebrate feeding on P. littoralis. The results indicated that (1) the grazing on F. vesiculosus depended on the availability of P. littoralis, (2) the photosynthetic activity of algae explained the best the variation in grazing rate and (3) the grazers are not likely to control the early outbreak of filamentous algae in the northern Baltic Sea by avoiding young and photosynthetically active algae. The likely mechanism behind the relationship is that the increased photosynthetic activity of macroalgae coincides with their higher resistance to herbivory.

Factors controlling long-term changes of the eutrophicated ecosystem of Pärnu Bay, Gulf of Riga

Phytoplankton, mesozooplankton, mysids and fish larvae were studied during 15–29 annual cycles measured weekly to monthly in Parnu Bay, the Gulf of Riga. The monthly variability of the biological data was related to temperature, ice conditions, salinity, influx of nutrients, the North Atlantic Oscillation (NAO) index, cloudiness and solar activity. Phytoplankton development was mainly a function of the NAO index. For the whole study period the abundance of zooplankton increased with increasing water temperature and solar activity. Significant correlations between phytoplankton and zooplankton densities were found until 1990. After the invasion of the predatory cladoceran Cercopagis pengoi in 1991, the zooplankton community was likely to be regulated by the introduced species rather than phytoplankton dynamics. The increased abundances of rotifers and copepods triggered the increase in mysid densities. The development of herring larvae was positively affected by the high density of copepods and rotifers but also by increased eutrophication. Until 1990 there was no significant relationship between the density of zooplankton and herring larvae. A negative relationship between the density of zooplankton and herring larvae in the 1990s suggests that the major shift in zooplankton community resulted in food limitation for herring larvae. The results indicated that (1) atmospheric processes in the northern Atlantic explain a large part of the interannual variation of the local phytoplankton stock, (2) trophic interactions control the development of pelagic communities at higher trophic levels, and (3) the introduction of an effective intermediate predator has repercussions for the whole pelagic food web in Parnu Bay.

Ecological consequence of the introduction of the polychaete Marenzelleria cf. viridis into a shallow-water biotope of the northern Baltic Sea

Effects of the North American polychaete Marenzelleria cf. viridis on a simple shallow-water benthic community of the northern Baltic Sea were studied in a field experiment combining natural densities of dominating macrofaunal species. The presence of M. cf. viridis increased benthic production (chlorophyll a) and reduced the survival of the native polychaete Nereis diversicolor. Adult Macoma balthica caused a significant mortality on M. cf. viridis whereas adult Cerastoderma glaucum had no effect on M. cf. viridis. We suggest that the competitive interactions between M. cf. viridis and M. balthica are a possible key factor determining the distribution pattern of M. cf. viridis in the Baltic Sea.

Ecosystem impacts of the widespread non-indigenous species in the Baltic Sea: literature survey evidences major limitations in knowledge

Invasion of non-indigenous species (NIS) is acknowledged as one of the most important external drivers affecting structure and functions of marine ecosystems globally. This paper offers literature-based analysis on the effects of the widespread (occurring in at least 50% of countries) and currently established NIS on ecosystem features in the Baltic Sea. It appears that out of the 18 NIS taxa studied, there are no published records on 28% of NIS for any of the seven impact categories investigated. When ecological impacts are known, laboratory experimental evidence dominates over field studies. Combined observations on impact strength, information type and confidence level suggest that the two benthic invertebrates, the polychaete Marenzelleria spp. and the zebra mussel Dreissenapolymorpha (Pallas 1771) exert the highest ecosystem impact. Despite continuously accumulating information on the NIS effects, however, the confidence of findings is still low. Thus, we still understand very little on both the direction and magnitude of the effects of even the most widespread NIS on the structure and dynamics of the Baltic Sea ecosystems. In order to increase reliability of such assessments, future research should be targeted towards spatially-explicit field surveys and experimenting of multitrophic systems, together with modelling of ecosystem impact.