Oxygen and salinity characteristics of predator–prey distributional overlaps shown by predatory Baltic cod during spawning

In the distributional overlap volume of Baltic cod Gadus morhua and its prey, studied in the Bornholm Basin in the southern Baltic Sea, only a fraction of the sprat Sprattus sprattus population vertically overlapped with the Baltic cod population. Sprat occurred in the intermediate water, in the halocline and in the bottom water, while herring Clupea harengus and Baltic cod occurred exclusively in the halocline and in the bottom water. Only parts of the sprat population were hence accessible for Baltic cod, and only a fraction of the sprat had access to the Baltic cod eggs below the halocline. Baltic cod–clupeid overlap volumes appeared to be determined by salinity stratification and oxygenation of the bottom water. Hydrography time series were used to estimate average habitat volumes and overlap from July to September in 1958–1999. In the 1999 survey spawning Baltic cod had greater ratios of empty stomachs and lower average rations than non-spawning Baltic cod. The average ration for Baltic cod caught within 11· 4 m from the bottom (demersal) did not differ from the average ration of Baltic cod caught in shallower waters (pelagic), because spawning and non-spawning Baltic cod in both strata were caught at equal rates. The diet of the Baltic cod caught demersally contained more benthic invertebrates, especially Saduria entomon, but Baltic cod caught pelagically also had fresh benthic food in their stomachs, indicating vertical migration of individual fish.     

Impact of the Alien Species <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> A. Agassiz, 1865 on Fish of the Caspian Sea

The data of long-term ecological studies (1999–2014) are used for assessing the impact of a new Caspian invader Mnemiopsis leidyi A. Agassiz, 1865 on marine and anadromous fish species. The introduction of the ctenophore and its massive development in the Caspian Sea have mainly affected planktophagous fish, especially the most abundant species anchovy sprat (Clupeonella engrauliformis Borodin, 1904) and sturgeons (Family Acipenseridae). Mnemiopsis leidyi is one of the main negative factors preventing the restoration of the populations of Caspian fishes, especially anchovy tyulka. The critical state of the resources of the spawning parts of sturgeon populations (Family Acipenseridae) has been reinforced as a result of the massive development of M. leidyi.     

Vertical distribution of eggs and larvae of the Baltic Sprat <Emphasis Type="Italic">Sprattus sprattus balticus</Emphasis> (Clupeidae) in relation to seasonal and diurnal variation

Vertical distribution of eggs and larvae of the Baltic sprat Sprattus sprattus balticus was studied. Diurnal variation was revealed in the distribution of eggs of the first stage of development and larvae with a length of more than 5 mm, which is related to dynamics of spawning and vertical migrations of sprat larvae. Seasonal variation in the distribution of eggs manifested itself in: (1) widening of the range of occurrence from the zone of halocline at the beginning of spawning (winter) to the entire water column in spring-summer and (2) emergence in the surface layer in July of all stages of egg development unlike the presence from May to June mainly of the first stage of development. It is suggested that the ability of early ontogenetic stages of sprat to maintain in the surface layer depended on the extent of development of vertical gradients of density in the thermocline.     

Fish eggs and larvae in the diet of herring Clupea harengus membras Linnaeus, 1758 and the sprat Sprattus sprattus balticus (Schneider, 1904) (Clupeidae) in the Southeastern Baltic Sea

The qualitative and quantitative composition of fish eggs and larvae in the stomach contents of Baltic herring and sprat was examined. The occurrence of fish eggs and larvae in the diet of herring was markedly higher, compared to sprat. Eggs of four and larvae of nine fish species were found in the herring stomachs. The prey species composition of the herring and sprat diets was related to the ichthyoplankton composition and to the degree of overlapping of the prey and predator distributions. Sprat eggs were the predominant prey in the herring diet in May and sprat larvae prevailed in August. The daily ration estimates indicate that the sprat larvae consumption by herring in August exceeded the sprat egg consumption in May by many times. Cod eggs in the herring stomachs were only found at the optimal oxygen content and salinity for the cod eggs in the near-bottom water layer.     

Timing and size of blooms of the ctenophore Mnemiopsis leidyi in relation to temperature in Narragansett Bay, RI

The ctenophore Mnemiopsis leidyi is at the northern extreme of its geographic range in Narragansett Bay, an estuary on the northeast coast of the United States. Blooms have typically been observed in late summer and fall according to records from 1950 to 1979. We document an expansion of the seasonal range of this important planktonic predator to include springtime blooms during the 1980s and 1990s. This shift to an earlier seasonal maximum is associated with increasing water temperature in Narragansett Bay. Temperatures in spring have risen, on average, 2 °C from 1950 to 1999 with warm years being associated with the positive phase of the North Atlantic Oscillation. During 1999, M. leidyi appeared earlier in spring and was more abundant than during any previous year for which records are available. Changes in the seasonal pattern and abundance of this predator are likely to have important effects on planktonic ecosystem dynamics of Narragansett Bay. These include reduction of zooplankton abundance in spring followed by increases in size and frequency of summer phytoplankton blooms. Earlier blooms of M. leidyi may also reduce survival of eggs and larvae of fish because, as in 1999, they coincide with the period of peak spawning.     

Ecology and long-term forecasting of sprat (<Emphasis Type="Italic">Sprattus sprattus balticus</Emphasis>) stock in the Baltic Sea: a review

In the brackish Baltic Sea situated in the transition zone between the Atlantic and Euro-Asiatic continental climate systems, the periods of high abundance of sprat of marine boreal origin coincide with a rich freshwater discharge, large water volume inhabitable by fish, rather high winter temperature and low salinity limiting the stock of its main predator—cod (Gadus morhua callarias). In the freshening periods an additional volume of water with acceptable oxygen and temperature conditions for sprat is formed in the Eastern (E) and, especially, the Northwest (NW) regions of the Baltic Sea. This allows extraordinary increase in sprat abundance/biomass. The conditions for sprat deteriorate during the period of active saline water inflows and colder winters, especially in the NW region. Following the decrease in the volume of water acceptable for the wintering of sprat as to temperature and/or oxygen concentration, some part of the stock may be forced to migrate southwards as in the Southwest (SW) region the conditions are the most stable. This may cause an extensive mixing of the stock components of various regions, hindering the differentiation of sprat regional units in the Baltic Sea. As no isolation of regional spawning concentrations has been found, beginning with 1989 the Baltic sprat has been assessed and managed as one stock unit, despite spatial differences in its morphological characters, growth rate etc. The alternation of periods of different regime-forming conditions (freshening or oceanization) is probably triggered by climate changes. This regularity has been exploited for the composition of long-term forecasts of qualitative changes in fish (incl. sprat) stocks in the Baltic Sea.     

Changes in biodiversity of phytoplankton,zooplankton, fishes and macrobenthos in the Southern Caspian Sea after the invasion of the ctenophore <Emphasis Type="Italic">Mnemiopsis Leidyi</Emphasis>

Monitoring for 6 years (2001–2006) showed that the population explosion of the alien ctenophore Mnemiopsis leidyi in the southern Caspian Sea coincided with a decline in the abundance and species number of mesozooplankton. While this decline appeared to have reduced the nourishment of sprat (also known as kilka), it seemed to have affected phytoplankton favorably mainly due to the decrease in grazing pressure. During 2001–2002, when M. leidyi abundance and biomass were at their highest levels, abundance of dinoflagellates and cyanophytes exceeded that of diatoms. Before the invasion (1996) and in some years after the invasion (2003, 2004 and 2006) diatom abundance was higher than the abundance of other groups. In September 2005, an unprecedented bloom of the toxic cyanophyte Nodularia sp. was observed in the southern Caspian Sea. Disappearance of edible zooplankton such as Eurytemora spp. was among the first changes observed after the expansion of M. leidyi in the area. Some changes in the macrobenthic fauna were also conspicuous after the increase of this ctenophore. While the biomass of some deposit feeders, such as the polychaete Nereis diversicolor and oligochaete species increased, benthic crustaceans decreased sharply in abundance during 2001–2003 and completely disappeared during 2004–2006. Iranian catches of kilka, the most abundant and widespread zooplanktivorous fish, decreased significantly in the southern Caspian Sea after 1999. Iranian landings of kilka dropped ~70% from 69,070 ± 20,270 t during 1995–2000 to 23,430 ± 12,240 t during 2001–2006, resulting in a loss of at least 125 million US dollars to the economy. There were also changes in the total catches of large predators such as the kutum and mullet, which mainly feed on kilka, between 1991 and 2006.     

Laboratory studies of predation by the ctenophore Mnemiopsis leidyi on the early stages in the life history of the bay anchovy, Anchoa mitchilli

The lobate ctenophore, Mnemiopsis leidyi, consumed eggs andlarvae of the bay anchovy, Anchoa mitchilli, in laboratory experiments.This ctenophore exhibited a type I functional response to increasesin egg densities without reaching saturation at high prey densities.Clearance rate increased with increasing experimental containervolume. There was a 3-fold increase in the volume of water clearedby 2.0–2.5-cm ctenophores and a >5-fold increase for4.5–5.0-cm ctenophores in 15–1 versus 100–200–1containers. Clearance rate was dependent on tenophore lengthbut was probably underestimated for the larger animals due tocontainer effects. The presence of various densities of alternateprey, Acartia hudsonica or Anemia sp. nauplii, in addition toAnchoa mitchilli eggs did not affect the clearance rates onthe eggs alone. Comparison of clearance rates of 2.0–2.5-cmctenophores on various ages of starved and fed bay anchovy larvaeindicated that predation may be higher on yolk-sac larvae thanon eggs but decreases as the larvae grow. After 3 days posthatch starved anchovy larvae become more vulnerable to predationthan fed larvae. The ctenophore, M.leidyi has the potentialto inflict substantial predation pressure on early stages inthe life history of bay anchovy. ¹ Present address: University of Maryland, Center for Environmentaland Estuarine Studies, Chesapeake Biological Laboratory, POBox 38, Solomons, MD 20688-0038, USA     

First evidence of spawning of eastern Baltic cod (Gadus morhua callarias) in the Belt Sea,the main spawning area of western Baltic cod (Gadus morhua L.)

Two cod stocks (western Baltic cod, WBC, and eastern Baltic cod, EBC) are managed in the Baltic Sea which is characterized by different main spawning areas and different main spawning periods. In this study we analyse the spatial and temporal occurrence of spawning individuals of both cod stocks in the main spawning grounds of the Baltic Sea based on eight microsatellite loci. Our results suggest that EBC (Gadus morhua callarias) has formed currently temporally stable, substantially homogeneous population not only in the Bornholm Sea (ICES SD: 25) but also in the Arkona Sea (ICES SD: 24). The presented analyses proved that EBC (G. m. callarias) can temporarily also spawn in the Belt Sea.     

The abundance and patterns of the egg and larval spatial distribution of Eastern Baltic cod Gadus morhua callarias (Gadidae) in 1931–1996

According to ichthyoplankton surveys of AtlantNIRO and published sources, the data on abundance of eggs and larvae of eastern Baltic cod Gadus morhua callarias at different spawning grounds of the Baltic Sea in 1931–1938, 1949–1956, and 1991–1996 are considered. We show that a similar type of spatial distribution of eggs and larvae of cod with localization of the greatest amount of its early ontogenetic stages in the Bornholm Basin in the southwest of the sea was observed in the 1930s and 1990s. The extension of the area of distribution was accompanied by an increase in the numbers of eggs and larvae of cod in all areas of the sea, and its decline was followed by a decrease in numbers, in the first turn, at northeastern spawning grounds in the Gotland Basin. The period of high numbers of eggs in 1949–1956 coincided with the century maximum of salinity of the near-bottom water layer.     

Somatic organic content of the ctenophores <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> (Ctenophora: Lobata) and <Emphasis Type="Italic">Beroe ovata</Emphasis> (Ctenophora: Beroida) in early ontogenetic stages

The organic matter content in the eggs and early larvae of the ctenophores Mnemiopsis leidyi and Beroe ovata from the Black Sea was determined using the adapted microtechnique of dichromate oxidation. The content of organic matter in the eggs of M. leidyi (0.25 μg/indiv.) was 5 times less than in B. ovata (1.28 ± 0.29 μg/indiv.). The somatic content of organic matter was 0.25 ± 0.09 μg/indiv. (25.1 ± 8.3 μg/mg, wet wt) for 2-day-old larvae of M. leidyi (0.2–0.3 mm in body length) and 1.37 ± 0.19 μg/indiv. (67.1 ± 5.7 μg/mg wet wt) for larvae of B. ovata (0.4 mm in body length). The specific organic content of larvae of both species steadily decreased with an increase in ctenophore body size and weight, approaching 3–4 μg/mg of wet weight for 2 mm specimens of M. leidyi and 3–5 μg/mg of wet weight for 6 mm B. ovata. The specific organic content of early larvae was 20–30 times higher than that of adult ctenophores. The results of this investigation could be useful in the evaluation of the energy budget for somatic growth and generative production in these species. Calculations indicate that with specific wet weight growth rates of 0.43/day for M. leidyi larvae and 0.29/day for B. ovata larvae, their true organic increases are respectively 30 and 38% less, i.e., no more than 0.31/day for the former and 0.18/day for the latter species.     

Spreading and physico-biological reproduction limitations of the invasive American comb jelly Mnemiopsis leidyi in the Baltic Sea

Hydrodynamic drift modeling was used to investigate the potential dispersion of Mnemiopsis leidyi from the Bornholm Basin in the Baltic Sea where it has been observed since 2007 further to the east and north. In the brackish surface layer dispersion is mainly driven by wind, while within the halocline dispersion is mainly controlled by the baroclinic flow field and bottom topography. Model runs showed that the natural spreading via deep water currents from the Bornholm Basin towards north and east is limited by topographic features and low advection velocities. Based on the information on ranges of salinity and temperature, which limit survival and reproduction of this ctenophore within the Baltic Sea, areas have been identified where the American comb jelly, M. leidyi could potentially survive and reproduce. While, we could show that M. leidyi might survive in vast areas of the northern Baltic Sea its reproduction is prevented by low salinity (<10 psu) and temperature (<12°C). Thus, due to the combined effect of low salinity and temperature, it is not probable that M. leidyi could establish permanent populations in the central or northern Baltic Sea. However, it seems that in the southern parts of the Baltic Sea environmental conditions are suitable for a successful reproduction of M. leidyi.     

Avoidance of hydrodynamically mixed environments by <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> (Ctenophora: Lobata) in open-sea populations from Patagonia,Argentina

Biomass distributions of the ctenophore, Mnemiopsis leidyi, were collected during a week-long survey program in a tidal front along the Patagonian shelf in December 1989. Average ctenophore biomass concentrations varied significantly along a north–south gradient and in stratified compared to unstratified waters. The relative vertical distribution of M. leidyi biomass appeared to be constrained by surface levels of vertical shear. Vertical distributions of ctenophore biomass were highly variable at low levels of vertical shear (<4 s⁻¹) at the surface, but at higher levels of surface vertical shear ctenophores occurred deeper in the water column where shear levels were lower. These results indicate that physical conditions are important factors influencing the distribution of M. leidyi along the Patagonian shelf during summer months.     

Predator-induced vertical behavior of a ctenophore

Although many studies have focused on Mnemiopsis leidyi predation, little is known about the role of this ctenophore as prey when abundant in native and invaded pelagic systems. We examined the response of the ctenophore M. leidyi to the predatory ctenophore Beroe ovata in an experiment in which the two species could potentially sense each other while being physically separated. On average, M. leidyi responded to the predator’s presence by increasing variability in swimming speeds and by lowering their vertical distribution. Such behavior may help explain field records of vertical migration, as well as stratified and near-bottom distributions of M. leidyi.     

Potential pathways of invasion and dispersal of <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> A. Agassiz 1865 in the Baltic Sea

The rapid spread of Mnemiopsis leidyi across the entire Baltic Sea after its first observation in 2006 gave rise to the question of its invasion pathway and the possible vector of its transport. To investigate pathways of M. leidyi invasion, the years 2005–2008 have been simulated by a three-dimensional coupled sea ice-ocean model of the Baltic Sea. In addition, a Lagrangian particle-tracking model has been utilized to test possible transport routes of this invader for 2006/2007. Based on the model, we exclude advection from the Kattegat as the main area of origin of M. leidyi and further spreading through the entire Baltic Sea. To explain the dispersion of M. leidyi in 2007 an earlier invasion already in 2005 is most probable. Alternatively, an invasion originating from main harbors with high ship traffic could also be a potential pathway. Drift simulations with drifter release in the main harbors are in good agreement with the observed distribution pattern of M. leidyi.     

Invaders Ctenophores Mnemiopsis leidyi (A. Agassiz) and Beroe ovata Mayer 1912, and Their Influence on the Pelagic Ecosystem of Northeastern Black Sea

We analyzed the state of pelagic ecosystem of northeastern Black Sea affected by invasion of a new settler Beroe ovata in 1999–2001. Development of B. ovata considerably decreased the population of another invader, Mnemiopsis leidyi, that deformed the Black Sea ecosystem for over a decade. Reduction of M. leidyi population limited its influence on the ecosystem and, consequently, we observed recovering of the main components of the Black Sea pelagic ecosystem—zooplankton and fish as well as their eggs and larvae. The relationships between annual and seasonal variability of the population development and size of the both invaders M. leidyi and B. ovata are discussed.     

Long-term prediction on Baltic fish stocks based on periodicity of solar activity

The methods presently in wide use for the assessment of marine ecosystems and fish stocks cannot provide the high-quality long-term prognoses urgently needed for improved management of marine ecosystems and their fishery resources. A novel method of forecasting the long-term qualitative composition of ecosystem and fish stocks in the Baltic Sea based on the periodicity and succession of the processes: extraterrestrial factors????climate changes????regime forming factors (chiefly salinity and temperature)????ecosystems????fish stocks was used in 1989 to predict the approximate year class abundances of cod (Gadus morhua callarias), sprat (Sprattus sprattus balticus), sea herring and gulf herring (Clupea harengus membras) in the Baltic Sea for the years 1989?C2008. A comparison with actual development up to 2008 show that this prognosis was confirmed to a considerable extent. The method based on periodical fluctuations of freshwater input and the thermal regime is described. It predicts a new regime shift in the 2020s to a higher salinity and more acceptable conditions for the organisms of marine origin.     

Seasonal changes and population dynamics of the ctenophore Mnemiopsis leidyi after its first year of invasion in the Kiel Fjord, Western Baltic Sea

We analyzed the seasonal variations of the ctenophore Mnemiopsis leidyi weekly collected since its first record in the western Baltic Sea in October 2006. The distribution pattern together with the seasonal dynamics and population outbreaks in late summer 2007 indicate recent successfully establishment of M. leidyi in this area. Seasonal changes showed two periods of high reproductive activity characterized by a population structure dominated by small size classes, followed by an increase of larger ones. These results further revealed that the bulk of the population remains in deep layers during the periods of low population density, whereas it appeared situated in upper layers during the proliferation of the species. We further emphasized the strength of the population outbreaks, which can reach abundances >10-fold higher in time periods shorter than a week. The predatory impact this species may have in pelagic ecosystems warns on the importance of its recent range of expansion.     

Uranium and thorium in muscle tissue of fish taken from the southern Baltic

The determination of U and Th was carried out on pooled samples of muscle tissue of cod (Gadus morhua), herring (Clupea harengus), sprat (Sprattus sprattus) and some other species of fish caught in 1981 in the southern Baltic. The levels of U obtained in the present study are generally within the ranges presented by other authors for fish from different aquatic regions of the world. The concentrations of U are similar to those found for Th in Baltic fish analysed. The concentration factors CF_U(Th), discrimination factors DF_U/Th and Th/U ratios (by weight) were calculated and discussed. These coefficients were also compared with values determined in other Baltic organisms. Transfer of U and Th along a food chain from potential prey (diet) to fish as predator (consumer) was quantified with a transfer factor.