Establishment of the Baikalian endemic amphipod Gmelinoides fasciatus Stebb. in Lake Ladoga

Gmelinoides fasciatus Stebb., a small amphipod from the Lake Baikal basin, was discovered in July 1988 in Lake Ladoga, the largest European lake. G. fasciatus likely invaded Lake Ladoga as a consequence of its intentional introduction, aimed at enhancing fish production, in some Karelian Isthmus lakes close to Lake Ladoga's western shore in the early 1970's. Benthos studies conducted in 1989 and 1990 revealed that G. fasciatus was well established in littoral communities along the western and northern shores of Lake Ladoga. G. fasciatus was the dominant species in these littoral communities and contributed over 70% of the macroinvertebrate biomass. The species was abundant in different macrophyte beds and stony littoral areas, both in heavily polluted and undisturbed sites. The maximum abundance and biomass of G. fasciatus was about 54,000 ind m^–2 and 160 g (wet wt.) m^–2. Negative impacts of G. fasciatus on native species have been observed. Studies are needed to identify the effects of G. fasciatus on the functioning of littoral communities and to predict and control the spread of this amphipod.     

Feeding selectivity,food assimilability and demand of the Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) in Lake Ladoga

Feeding selectivity and assimilability of the Baikal invader Gmelinoides fasciatus (Stebbing, 1899) in Lake Ladoga at feeding by various types of plant and animal food have been studied experimentally. The food demand has been assessed at different assimilation levels. Freshwater shrimp G. fasciatus has been shown to have distinct selective capacity with respect to different kinds of plant and animal food.     

Population characteristics and life cycle of the Lake Baikal invader <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis> (Stebbing, 1899) (Crustacea: Amphipoda in Lake Ladoga

Dynamics of the size-age structure and sexual structure of the population and characteristics of reproduction and life cycle have been studied in the Baikal invader Gmelinoiudes fasciatus (Stebbing, 1899) in Lake Ladoga. The studied characteristics display both similar features with and differences from populations of this species in other waterbodies. It is found that the size and age structure of the G. fasciatus population changes in the circannian aspect under the influence of complex of environmental factors. The sexual structure of the population is dynamic, but a sex ratio of approximately 1: 1 is maintained in the circannian cycle. Six principal periods of hatching are recognized. The main factor determining the timing of the start and termination of the reproduction period in G. fasciatus is temperature. The weight of eggs oviposited by the female over the whole life cycle (about 21.5 cal in the energy equivalent) amounts to about 60% of the female body weight.     

Distribution,population structure and salinity tolerance of the invasive amphipod Gmelinoides fasciatus (Stebbing) in the Neva Estuary (Gulf of Finland,Baltic Sea)

In the early 1970s, the Baikalian amphipod Gmelinoides fasciatus (Stebbing) was intentionally introduced into several lakes in the Gulf of Finland basin in order to enhance fish production. By 1996, G. fasciatus successfully colonized the littoral zone of Lake Ladoga and, via the Neva River, invaded the Neva Bay, the freshwater part of the Neva Estuary. In 1999, G. fasciatus was first registered in the inner Neva Estuary, the very first record of the Baikalian amphipod in brackish waters of the Baltic Sea. Distribution, abundance, reproduction and population structure of G. fasciatus in the Neva Estuary were studied during 1998–2000. In some locations of the Neva Estuary, maximum densities of G. fasciatus reached 3500 ind. m⁻². In general, density and biomass of G. fasciatus in the freshwater part of the Neva Estuary were higher (around 1.5 fold) than in the brackish-water part. Fecundity of this amphipod averaged 10–20 eggs per female, depending on body size of females and season. In order to assess the possibility of further spread of G. fasciatus in the Baltic Sea, the salinity tolerance of this species was determined in a series of laboratory experiments. Our results showed that the invasive amphipod G. fasciatus is potentially able to colonize shallow coastal habitats of, for example, the Gulf of Bothnia, Gulf of Riga and other parts of the Baltic Sea with water salinities ranging from 1 to 5 psu.

     

Heat shock proteins in the mechanisms of stress adaptation in Baikal amphipods and Palaearctic <Emphasis Type="Italic">Gammarus lacustris</Emphasis> Sars II. Small HSP family

The role of low-molecular-weight (small) heat shock proteins (sHSPs) in the mechanisms of thermal and toxic resistance in fresh-water organisms was investigated. The four endemic species of Lake Baikal: Gmelinoides fasciatus (Stebb.), Eulimnogammarus cyaneus (Dyb.), E. vittatus (Dyb.), and Ommatogammarus flavus (Dyb.), and Gammarus lacustris Sars, a representative of the Palaearctic fauna, were studied. The effect of the temperature factor was evaluated by exposure of amphipods to temperatures of 20, 25, and 30°C, and the effect of the toxic factor, by exposure to cadmium chloride solutions with concentrations of 50, 10, 5, 0.5, and 0.05 mg/l. All the species showed a common trend of increasing content of sHSPs; species-specific features were observed in the synthesis of the proteins. It was concluded that sHSPs participate in the mechanisms of thermal and toxic resistance in the investigated amphipod species.     

The Mechanisms for Genetic Diversity of Baikal Endemic Amphipod <Emphasis Type="Italic">Gmelinoides fasciatus</Emphasis>: Relationships between the Population Processes and Paleoclimatic History of the Lake

Amphipods of the species Gmelinoides fasciatus are one of the dominant species in the littoral zone community of Lake Baikal. Earlier, on the basis of the analysis of the mitochondrial COX1 gene, four isolated populations within the species range were identified, between which there were no distinct geographical barriers. In the present study, a phylogenetic reconstruction of the evolutionary history of G. fasciatus with the molecular clock hypothesis is carried out. The population divergence time is dated back to the onset of global climate changes on the Pleistocene–Pliocene boundary at 1.8–2.3 Ma. As a result, the estimated substitution rate in the COX1 gene for G. fasciatus was 1.2–2.4% of substitutions per Myr, which corresponded to the values determined for other groups of invertebrates. It was demonstrated that geographical barriers that formed the populations of G. fasciatus periodically appeared and disappeared in the past, resulting from climate changes associated with the periods of global cooling and warming.     

Heat shock proteins in the mechanisms of stress adaptation in Baikal amphipods and palearctic <Emphasis Type="Italic">Gammarus lacustris</Emphasis> Sars. I. HSP70 family

The participation of HSP70 family proteins in the mechanisms of stress adaptation under thermal and toxic stress in Baikal endemic and palearctic organisms was investigated. The four Lake Baikal endemic species Gmelinoides fasciatus (Stebb.), Eulimnogammarus cyaneus (Dyb.), E. vittatus (Dyb.), Ommatogammarus fl avus (Dyb.) were compared with the palearctic fauna representative Gammarus lacustris Sars. The type of synthesis of HSP70 family heat shock proteins under thermal (exposure at a temperature of 20, 25, and 30°C) and toxic (exposure in CdCl₂ solutions with the concentrations 50, 10, 5, 0.5, and 0.05 mg/l) stress was determined. All species under investigation were shown to have a common trend towards HSP70 content increase; therewith some species-specific characteristics of the type of synthesis of the protein under investigation were observed. The conclusion has been made on HSP70 participation in the mechanisms of thermal and toxic resistance in the studied amphipod species.     

A transcriptome-wide assessment of differentially expressed genes among two highly divergent,yet sympatric,lineages of the freshwater Atyid shrimp, <Emphasis Type="Italic">Paratya australiensis</Emphasis>

The current study evaluated the possible toxic effects of the water-soluble fraction of crude oil on the general cellular stress-response mechanisms of two dominant representatives of Lake Baikal’s littoral community, the endemic amphipod species Eulimnogammarus verrucosus and E. cyaneus. The acute toxicity effects on the cellular stress-response mechanisms of amphipods were studied in the laboratory by exposing amphipods in water from Lake Baikal to addition of a water-soluble fraction of crude oil at concentrations considered safe for the aquatic environment. The present study found that even short-term exposure to a water-soluble fraction of crude oil at concentration of 50 µg/L, established as the threshold limit for fishery and aquaculture water reservoirs in the Russian Federation, directly affected the general stress-response markers HSP70 and lipid peroxidation and significantly changed the activity of antioxidant enzymes in both studied species. This result confirms the high sensitivity of Baikal endemics to crude oil. Thus, it also indicates that established standards and threshold limit values of oil concentrations estimated for ecological monitoring of general water reservoirs cannot be applied directly to the unique Lake Baikal ecosystem.     

Special traits of growth and feeding of the stone sculpin <Emphasis Type="Italic">Paracottus knerii</Emphasis> (Cottidae) in water bodies of different types

Growth and feeding of the stone sculpin Paracottus knerii (Cottidae) is investigated in Water Bodies of Different Types: Baikal (the littoral of its southern part), the Angara River, and the mountainous Sobolinoe Lake (the southern Baikal basin). The highest growth rate is characteristic of the South-Baikalian population, followed by the Angara population. The stone sculpin from Sobolinoe Lake is slow-growing. The growth rate of specimens of the latter is by 1.5–2 times lower than in the two aforementioned populations due to inferior feeding conditions of this water body. The size-age changes in the food spectrum of all investigated populations are consecutive change of all principal components and increase in size of the consumed prey     

Lake Superior: an invasion coldspot?

Lake Superior receives a disproportionate number of ballast water discharges from transoceanic ships operating on the Laurentian Great Lakes. Although this provides dispersal opportunities for nonindigenous species (NIS), relatively few NIS were initially discovered in this lake prior to being recorded elsewhere in the basin. A lack of NIS records from this lake may be an artefact of sampling bias. We tested this hypothesis by sampling benthos and plankton from littoral and deepwater habitats around the perimeter of Lake Superior during June and August 2001. Using morphological analysis techniques, we identified a total of 230 invertebrate taxa representing planktonic, benthic and nektonic lifestyles. Five species with invasion histories in the lower Great Lakes, the bivalves Sphaerium corneum, Pisidium amnicum and P. moitessierianum, gastropod Potamopyrgus antipodarum and amphipod Echinogammarus ischnus, were identified for the first time in Lake Superior. In addition, records of expanded distributions within this lake are presented for the amphipod Gammarus fasciatus and oligochaetes Ripistes parasita and Vejdovskyella intermedia. Recently introduced NIS in Lake Superior were found near international ports, implicating shipping as the vector of their introduction. Intrinsic physical-chemical aspects of Lake Superior may account for the scarcity of NIS in this lake as compared to the lower Great Lakes.     

Interspecific differences in drift behaviour between the native <Emphasis Type="Italic">Gammarus pulex</Emphasis> and the exotic <Emphasis Type="Italic">Gammarus roeseli</Emphasis> and possible implications for the invader’s success

“Drifting” is known to subject aquatic invertebrates to intense predation by drift feeding fish. Consequently, interspecific variations in drifting behaviour could lead to differences in predation pressure between coexisting prey species. Predation being an important factor determining the success of invaders, differences in drift patterns could advantage either native or exotic invertebrates through differential predation by native fish predators. The exotic freshwater amphipod (Gammarus roeseli) has now largely colonized Western Europe where it is often found in sympatry with a native species (Gammarus pulex). Here we documented interspecific differences in drifting behaviour that might have favored the invader’s success through differential predation. Benthic and drifting amphipods were sampled three times at the same site to compare the proportion of each species within and between sample types (benthos or drift) across time. Compared with the benthos, where the invader (G. roeseli) was significantly less abundant than the native (G. pulex), G. roeseli was proportionally overrepresented in the drift but displayed a very different drifting pattern. While G. pulex drift rates remained roughly constant over a 24 h period, G. roeseli showed a marked diel periodicity with low diurnal and high nocturnal drift rates. Such drifting behaviour could procure this species with a competitive advantage regarding predation as most drift feeding fish are diurnal. As a result, the native appears more disadvantaged with respect to drift. This may partly explain the ability of G. roeseli to coexist with G. pulex in a habitat more suitable to the native.     

Investigation of distribution,species composition,and degree of resistance to antibiotics of the bacteria of the <Emphasis Type="Italic">Enterococcus</Emphasis> genus in Lake Baikal

The paper presents the result of investigation of species composition, distribution and degree of resistance to antibiotics of the Enterococcus bacteria in Lake Baikal. It is shown that they live in littoral zones of the lake. In the deep-water part of the lake the studied microorganisms are not found. We isolated 120 strains identified as E. faecium, E. avium, E. faecalis, E. mundii, E. hirae, E. durans, E. gallinarum. On the whole, the strains of enterococcus isolated from Lake Baikal are characterized as antibiotic-sensitive. Nevertheless, the antibiotic-resistant strains of enterococci are found in the studied collection, including those with typical intermediate level of resistance to vancomycin.     

The role of the heat shock proteins (HSP70 and sHSP) in the thermotolerance of freshwater amphipods from contrasting habitats

The aim of this study was to clarify the significance of HSP70 and sHSP for thermotolerance in freshwater amphipods. We compared four amphipod species from different freshwater habitats and biogeographical regions (Central Europe vs. Lake Baikal). Test individuals were exposed to thermal stress generated by a water temperature of 25 °C. The thermotolerance of the species, determined by median lethal time (LT50), followed in decreasing order by Gmelinoides fasciatus, Echinogammarus berilloni, Gammarus pulex, Eulimnogammarus verrucosus. HSP70 and sHSP base level concentrations for the species were determined at control (i.e. non-stress) conditions. For HSP70, the base levels were positively correlated to the species' thermotolerances. For sHSP, however, only thermotolerant G. fasciatus showed a high level. Thermal stress at 25 °C water temperature caused a deferred onset of HSP70 and sHSP expression followed by a subsequent offset, delineating a unimodal response curve. The time lag to the expression onset of HSP70 was shorter in the thermosensitive species, compared to thermotolerant ones. Conversely, the time span until the maximum level of HSP70 was variable, not showing a dependence on the thermotolerance properties of the species. The peak concentration in G. pulex was distinctly higher than in the other species, whereas E. verrucosus did not develop a well-defined response maximum at all. In sHSP, the temporal pattern of expression was even more variable than in HSP70. However, the thermosensitive species E. verrucosus showed a time lag of expression onset significantly shorter than the other species and thermotolerant G. fasciatus developed the most pronounced response maximum. Basing on these results, the cellular response to thermal stress in amphipods is more consistently reflected by HSP70, compared to sHSP.     

Two new species of Hyalella from Lake Titicaca,and redescriptions of four others in the genus (Crustacea: Amphipoda)

The amphipod genus Hyalella has its highest diversity in the Andean Lake Titicaca. This genus is the only epigean amphipod present in South America. Eleven endemic and one non-endemic species are known from Lake Titicaca. An additional endemic species, Hyalella nefrens n. sp. and one non-endemic species Hyalella tiwanaku n. sp. are described here. Hyalella cuprea(Faxon, 1876), Hyalella latimanus (Faxon, 1876), Hyalella montforti Chevreux, 1907, and Hyalella neveulemairei Chevreux, 1904 are redescribed. The high diversity of Hyalella in the lake is not well understood, and it has been compared with the spectacular diversity of Lake Baikal.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Distribution and summer diapause of Canthocamptus staphylinus (Jurine) (Copepoda: Harpacticoida) in Lake Ladoga

The spatial distribution of Canthocamptus staphylinus (Jurine) in Lake Ladoga was examined. C. staphylinus is one of the most numerous species of harpacticoids in the lake. It can dwell in almost all littoral biotopes. In profundal biotopes strongly exposed to anthropogenic influence (pollution and eutrophication) only this species of harpacticoid was found. A peculiarity of C. staphylinus biology is that in summer individuals are in the resting stage as cysts. In summer active individuals were found in littoral habitats at a water temperature of 20 °C, but their number was small in comparison with cold periods of the year. In the littoral zone the relative number of individuals in cysts was clearly related to water temperature. Twelve degrees temperature appears to be critical for development of littoral populations of C. staphylinus.     

Feeding of the stone sculpin Paracottus knerii (Cottidae) in the littoral of southern Baikal

The food spectrum and seasonal dynamics of feeding of the stone sculpin Paracottus knerii in the littoral of southern Baikal is investigated. Altogether, 29 species of invertebrates are recorded. All year round, the bulk of food, similarly to other areas of the lake, consisted of amphipods; other groups of zoobenthos were represented insignificantly. The significance of particular amphipod species in the food of fish depended on their ecological and morphofunctional traits. The seasonal changes in the species composition and the ratio of food items in the food depended on life cycles of fish and on the migrations of zoobenthic organisms.     

Environmental heterogeneity limits the local dominance of an invasive freshwater crustacean

The impacts of an exotic species may vary along environmental gradients. We tested the hypothesis that site-specific heterogeneity in the physico–chemical environment alters the dominance of two competing amphipods: Gammarus fasciatus, the dominant native species in the upper St. Lawrence River, and Echinogammarus ischnus, a euryhaline Ponto-Caspian species that invaded the river in the late 1990s. E. ischnus has replaced G. fasciatus as the dominant amphipod at some sites, while remaining inferior at other sites—even several years after invasion. We tested the effect of water chemistry (conductivity) on the outcome of interactions between these two species in the laboratory and in the river in 2006 and 2007. Field experiments involved transplanting both species to different sites along a gradient of conductivity within the river, whereas the laboratory experiments examined interspecific predation in water collected from these sites. The laboratory experiments revealed that these species are mutual predators and their rate of predation on each other varies with conductivity; E. ischnus is the dominant predator at higher conductivity, while G. fasciatus dominates at lower conductivity. The field experiments showed interannual variation, but supported the lab results at high conductivity in 2006 and at low conductivity in 2007. E. ischnus was more sensitive than G. fasciatus to variation in conductivity, which apparently mediates this invader’s ability to replace or dominate the native species in a heterogeneous environment.     

The consortium of the sponge Lubomirskia baicalensis in Lake Baikal,East Siberia

The spatial distribution of the fauna associated with a branched sponge, Lubomirskia baicalensis, endemic of Lake Baikal has been quantitatively studied. The biomass and numbers of three amphipod species which inhabit the sponge correlate (linearly or non-linearly) with the weight of the sponge.     

Seasonal dynamics of growth and feeding of big-headed sculpin <Emphasis Type="Italic">Batrachocottus baicalensis</Emphasis> (Cottidae) in the Cape Berezovyi area (southern Baikal)

The seasonal features of the growth and feeding of one of the mass fish species of the littoral zone of Lake Baikal, big-headed sculpin Batrachocottus baicalensis, are investigated in the study area near the Cape Berezovyi (southern Baikal). It is found that the greatest increase of the linear parameters of the individuals of all age classes occurs in autumn, in the middle of the feeding period. Twenty-nine food objects, i.e., invertebrates and fishes, have been noted in the feeding spectrum of big-headed sculpin. It is found that the amphipods comprise the basic food year round. The transition of big-headed sculpin to predominant consumption of fish food is noted in the reproduction periods of other sculpins in the shallows. In general, the seasonal changing in species composition and food object ratio in the food are due to the fish life cycles, migrations of amphipods of different species, and, to a lesser extent, changing numbers of different groups of zoobenthos.