Invasions of alien gammarid species and retreat of natives in the Vistula Lagoon (Baltic Sea, Poland)

During the last decades of the twentieth century, the alien gammarid species Gammarus tigrinus, Dikerogammarus haemobaphes, Pontogammarus robustoides and Obesogammarus crassus invaded the lower Vistula River and its deltaic, partly brackish regions. In brackish waters of the Vistula Lagoon the native Atlantic-boreal species Gammarus zaddachi and Gammarus duebeni have been replaced or at least outnumbered by the aliens. As compared to our earlier studies, through the years 1998–2004 we could observe nearly total decline of the native gammarid populations along the coasts of the Lagoon, and overdomination of the North-American G. tigrinus in most places. Possible reasons for the observed phenomena are e.g. increasing pollution and eutrophication of the Lagoon accompanied by competition between the native and the alien species.     

There is more than one ‘killer shrimp’: trophic positions and predatory abilities of invasive amphipods of Ponto‐Caspian origin

1. Biological invasions are regarded as one of the greatest threats to biological diversity. One of the macroinvertebrate groups with the largest number of invasive species in fresh water are gammarid amphipods. Their omnivorous (including predatory) feeding behaviour may facilitate their spread and establishment in new areas. 2. Dikerogammarus villosus, the ‘killer shrimp’, is a well‐known example of a Ponto‐Caspian gammarid that is a very effective predator and successful coloniser in Europe. There are, however, other invasive Ponto‐Caspian amphipods, which have spread successfully in Northern, Central and Western Europe. Our aim here was to test whether two of such invaders (Pontogammarus robustoides and Dikerogammarus haemobaphes) are also more predacious than a native species (Gammarus fossarum). 3. Stable isotope analysis (δ¹⁵N and δ¹³C) of Ponto‐Caspian amphipods coexisting in a reservoir demonstrated that the trophic positions of P. robustoides and D. haemobaphes were similar to that of D. villosus. Echinogammarus ischnus and Chelicorophium curvispinum occupied the lowest position in the food web, while the native Gammarus fossarum (collected from another waterbody) had an intermediate trophic position. 4. Stomach content analysis of P. robustoides, D. haemobaphes and G. fossarum collected in the field, as well as laboratory feeding experiments, was used to compare diet and feeding preferences among the two invasive and one native species. All three species were omnivorous and predacious. However, the two invasive species (P. robustoides and D. haemobaphes) were more effective predators than G. fossarum and showed a clear preference for animal prey and tissue. 5. Pontogammarus robustoides and D. haemobaphes may, like D. villosus, also be called ‘killer shrimps’ and could have a similar impact as invaders of European freshwater and brackish waterbodies.     

How to be an invasive gammarid (Amphipoda: Gammaroidea)–comparison of life history traits

About six life history and two ecological traits of gammarid species occurring in Central European waters were compared in order to identify the characters of successful invader. The species were (1) natives: Gammarus fossarum, G. pulex, G. lacustris, G. varsoviensis, G. balcanicus, G. leopoliensis, G. roeselii, and (2) aliens: Gammarus tigrinus, Chaetogammarus ischnus (=Echinogammarus ischnus), Pontogammarus robustoides, Obesogammarus crassus, Dikerogammarus haemobaphes, D. villosus. Generally the alien species were characterised by a combination of large brood size, high partial fecundity, early maturation and by appearance of higher number of generations per year. Also, these species presented higher tolerance towards severe environmental conditions, i.e. elevated salinity and human degradation of the environment. The above features seem to facilitate the colonisation of new areas and competition with native species -- a phenomenon that has been currently observed in various parts of Europe.     

Species Composition and Distribution of Invasive Ponto‐Caspian Amphipods in the Off‐Channel Microhabitats of a Temperate,Lowland Dam Reservoir

The Włocławek Dam Reservoir located on the lower Vistula River (central Poland) is part of the central corridor used by Ponto‐Caspian species to migrate in Europe. It provides a number of habitats suitable for sustaining populations of several non‐indigenous taxa. Four Ponto‐Caspian amphipod species were recorded in the reservoir: Chaetogammarus ischnus, Chelicorophium curvispinum, Dikerogammarus haemobaphes and Pontogammarus robustoides. We found significant differences in amphipod densities and species composition among various microhabitats in off‐channel areas. P. robustoides was the only amphipod species that occurred on very shallow (<1 m) sandy bottom near the shore. It inhabited also other sites but its share in the total number of amphipods and abundance decreased with the distance from the shore. Furthermore, at sites more distant from shore its affinity for plant substratum was higher. D. haemobaphes, C. ischnus and C. curvispinum clearly preferred sites distant from shore, overgrown by macrophytes or covered by mussel shells. Furthermore, the abundance of C. ischnus was negatively correlated with the presence of P. robustoides. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Population dynamics of alien gammarid species in the River Odra estuary

During the period of 1988–2000, three alien gammarid species, i.e. Gammarus tigrinus Sexton, 1939, Pontogammarus robustoides G. O. Sars, 1894, and Dikerogammarus haemobaphes (Eichwald, 1841) were recorded in the Odra estuary. This study, the first record of G. tigrinus and P. robustoides in Poland, focuses on the distribution, abundance, and biomass as well as on population structure of the three new species found in the coastal zone of the estuary. G. tigrinus and P. robustoides were first sampled in the Szczecin Lagoon (mid-part of the estuary) in 1988. The densities of both species were high from the time they were first recorded and exceeded 10 000 ind. m^–2, while the biomass was 25 and 140 g m^–2, respectively. G.tigrinus occurred at higher densities in northern and eastern parts of the Lagoon, while P. robustoides was particularly abundant in the south-eastern part, most heavily affected by River Odra discharge. In 1999, the third gammarid, D. haemobaphes, was recorded in the southern part of the estuary. Both in 1999 and 2000, the species was found in qualitative samples (dredge collections of Dreissena polymorpha) taken from the western branch of the river. As estimated from quantitative samples, densities and biomass in the eastern branch were rather low and did not exceed 150 ind. m^–2 and 2.7 g m^–2, respectively. The Szczecin Lagoon population of G. tigrinus produces at least two generations within a year: a summer one and an overwintering one. The latter is dominated by older and larger specimens. The species may reproduce in the Lagoon from April until November.     

Length-weight relationships of Ponto-Caspian gammarids that have overcome the salinity barrier of the southern Baltic Sea coastal waters

Gammarids from the Caspian complex have invaded many European waters along the rivers and canals of the inland migration corridors. The species examined in this work are well known as invaders of European freshwater environments, so the colonization of brackish habitats is a phenomenon inviting more detailed investigation. The aim of this study was to determine the condition of the Ponto-Caspian gammarids Pontogammarus robustoides (G.O. Sars, 1894), Obesogammarus crassus (G.O. Sars, 1894), Dikerogammarus haemobaphes (Eichwald, 1841) and Dikerogammarus villosus (Sowinsky, 1894) as expressed by the relationship between total length and the wet weight of specimens in the brackish waters of the Gulf of Gdansk (Poland). The relationships can be regarded as responses to a newly expanded habitat after they overcome the salinity barrier of the southern Baltic Sea coastal waters. All these Ponto-Caspian gammarids demonstrated an increase in weight with increasing total length: P. robustoides (b = 2.852), O. crassus (b = 3.3477), D. haemobaphes (b = 3.7855) and D. villosus (b = 2.6917). The results are an indicator of the relatively good condition of the organisms and indicate that the brackish environment of the Gulf of Gdansk affords them excellent possibilities for growth.     

Effect of salinity on the distribution of Ponto-Caspian gammarids in a non-native area – environmental and experimental study

The native area of gammarids from the so-called ‘Caspian complex’, Pontogammarus robustoides (G.O. Sars, 1894), Obesogammarus crassus (G.O. Sars, 1894), Dikerogammarus haemobaphes (Eichwald, 1841) and D. villosus (Sowinsky, 1894), is associated with brackish waters. Over the last several decades they have colonized the European inland waters and part of the brackish Baltic Sea. It is believed that anthropogenic increase in the salinity of inland waters facilitated their expansion. However, the influence of salinity on the dispersal of gammarid species outside their native area is not fully understood. We tested the hypothesis that salinity was a major factor in determining distribution, based on the abundance of Gammaridae in three coastal areas of low salinity (brackish Baltic), i.e. 0.3, 3.4 and 7.3 PSU, successfully inhabited by them. Additionally, for the first time, the effect of water salinity on the osmoregulatory capacity of O. crassus was examined under laboratory conditions, for the salinities given above. The experiments showed that similarly as in the case of other Caspian complex species, salinity values of about 7 PSU create better conditions for osmoregulation in O. crassus than lower salinities (i.e. 0.3 and 3.4 PSU). In the environmental part of the study, we observed that only D. villosus achieved a significantly higher abundance in the area of 7.3 PSU. Thus, we concluded that in the range of 0.3–7.3 PSU, salinity is not a key factor governing the distribution of Ponto-Caspian gammarids.     

Exotic and indigenous freshwater gammarid species: physiological tolerance to water temperature in relation to ionic content of the water

Bioinvasions by closely related species often lead to niche competition between exotic and indigenous species. The outcome of this competition is partly determined by differences in physiological tolerance of the competing species to the environmental conditions of the colonised habitat. Physiological tolerance of the invading gammarid species Gammarus tigrinus, Echinogammarus ischnus and Dikerogammarus villosus and the indigenous gammarid species Gammarus pulex, Gammarus roeseli and Gammarus fossarum from Dutch waters was studied in the laboratory by comparing their pleopod beats at rest at different water temperatures, which reflect the gammarid's oxygen consumption. Pleopod beat frequencies increased from a minimum ventilatory activity of 0 beats per minute at 1 °C to maximum activity of up to 300 beats per minute at temperatures between 25 °C and 35 °C. At the state of maximum activity, a further increase in temperature was followed by a strong decrease in pleopod beat frequency, indicating acute stress, and subsequently mortality. Frequency response patterns of invading and indigenous gammarids were found to be highly similar, indicating a wide tolerance to temperature for all species. The tolerance of D. villosus, however, was reduced in brook water, indicating a lower competitive ability in relatively ion-poor water. G. tigrinus survived at higher temperatures in the more ion-rich, polluted waters than the indigenous gammarids, indicating a wider physiological tolerance and thus a higher competitive ability in these waters.     

Is salinity an obstacle for biological invasions?

Invasions of freshwater habitats by marine and brackish species have become more frequent in recent years with many of those species originating from the Ponto‐Caspian region. Populations of Ponto‐Caspian species have successfully established in the North and Baltic Seas and their adjoining rivers, as well as in the Great Lakes–St. Lawrence River region. To determine if Ponto‐Caspian taxa more readily acclimatize to and colonize diverse salinity habitats than taxa from other regions, we conducted laboratory experiments on 22 populations of eight gammarid species native to the Ponto‐Caspian, Northern European and Great Lakes–St. Lawrence River regions. In addition, we conducted a literature search to survey salinity ranges of these species worldwide. Finally, to explore evolutionary relationships among examined species and their populations, we sequenced the mitochondrial cytochrome c oxidase subunit I gene (COI) from individuals used for our experiments. Our study revealed that all tested populations tolerate wide ranges of salinity, however, different patterns arose among species from different regions. Ponto‐Caspian taxa showed lower mortality in fresh water, while Northern European taxa showed lower mortality in fully marine conditions. Genetic analyses showed evolutionary divergence among species from different regions. Due to the geological history of the two regions, as well as high tolerance of Ponto‐Caspian species to fresh water, whereas Northern European species are more tolerant of fully marine conditions, we suggest that species originating from the Ponto‐Caspian and Northern European regions may be adapted to freshwater and marine environments, respectively. Consequently, the perception that Ponto‐Caspian species are more successful colonizers might be biased by the fact that areas with highest introduction frequency of NIS (i.e., shipping ports) are environmentally variable habitats which often include freshwater conditions that cannot be tolerated by euryhaline taxa of marine origin.     

Intraguild predation and species exclusions in amphipods: the interaction of behaviour, physiology and environment

1. Data from field surveys, laboratory experiments and computer simulations of community dynamics revealed that a novel interaction among intraguild predation, physiological adaptation and environment may explain the complex distributions of two putatively competing aquatic amphipods. 2. Gammarus pulex and G. tigrinus both thrive in fresh and oligohaline waters in western Europe. However, the native European G. pulex excludes the invading North American G. tigrinus from freshwaters of relatively low conductivity, whereas the reverse occurs at higher conductivities. Additionally, there is much spatio-temporal fluctuation in the patterns of coexistence of these species. 3. Laboratory experiments in The Netherlands and Ireland revealed that mutual predation of moulting individuals occurred frequently between these species. However, predation frequencies were differentially in favour of G. pulex under the ionic conditions to which this species is physiologically adapted (freshwater). On the other hand, predation was not differential under the ionic conditions to which G. tigrinus is physiologically adapted (oligohaline water). 4. A mathematical model, which extends the logistic equation to include mutual intraguild predation, simulated interactions over a range of values of relevant population parameters. This indicated that G. pulex would be excluded when balanced instantaneous rates of mutual predation were combined with the known greater reproductive output of G. tigrinus. However, this reproductive advantage is overcome by any relatively small bias in the instantaneous rate of predation favouring G. pulex, leading to the exclusion of G. tigrinus. This occurs even when the reproductive advantage to G. tigrinus is relatively large. Moreover, the model generated ‘switches’ in species dominance that are determined by the relative values of reproductive rate and mutual predation. The time taken to ‘switch’ may explain the transient periods of apparent coexistence of these species observed in the field. 5. The complex community dynamics of such species may thus be understood in terms of variation in the intensity of species interactions mediated by behavioural, physiological and environmental factors.     

Reproductive biology of Dikerogammarus haemobaphes: an invasive gammarid (Crustacea: Amphipoda) colonizing running waters in Central Europe

Dikerogammarus haemobaphes is a Ponto-Caspian gammarid that has invaded vast areas in Central and Western Europe. Our paper is a first presentation of its life history features in an invaded region. The study was conducted in the Vistula River in Poland from autumn 2003 to autumn 2005 in two sites differing in hydrological conditions with one being water reservoir. The results showed that the reproductive period lasted 8 months from April till October in both sites. Three generations per 1 year were observed: autumn (overwintering), spring and summer. Ten cohorts per year were distinguished. The individuals from the reservoir were much bigger than those from the other site. The fecundity of those specimens was also higher and they laid 52 eggs per clutch in average in comparison with 37 eggs in the river itself. The strong relationship between the number of embryos (in developmental stage 2) per clutch and the length of females was noticed. The overall mean egg size of stage 2 of D. haemobaphes was 0.430 ± 0.029 mm which is smaller than noted for native species such as Gammarus fossarum. A potentially high reproductive capacity, comparatively small eggs, short time of eggs’ development, fast reaching sexual maturation, short life span, tolerance to a wide range of environmental conditions, all promote the invasion of this Ponto-Caspian gammarid in freshwater ecosystems of the temperate climate zone.     

Interference competition between alien invasive gammaridean species

The relative abundances of gammaridean species in the river Rhine have profoundly changed since the invasion of Dikerogammarus villosus in 1994/1995. This study tested whether these changes in gammaridean dominance could have been determined by interspecific competition and unequal mortality, for example by intraguild predation (IGP). Single and two species tests have been carried out in aquariums provided with all substrata present in the main channel of the Rhine. Changes in substratum choice, increased swimming activity and increased mortality of a species were used as indicators of interspecific competition during interaction between gammaridean species. Interspecific competition and mortality between the most abundant invasive gammaridean species in the Rhine, viz. Gammarus tigrinus, Echinogammarus ischnus and Dikerogammarus villosus were tested. In single-species experiments, G. tigrinus and D. villosus showed similar preferences for a stony substratum, whereas E. ischnus mostly occupied the water column. The two-species aquarium experiments indicated direct interference competition for substratum and unequal mortality between G. tigrinus and D. villosus, with D. villosus being the stronger competitor. Competitive stress was influenced by population density, was size-dependent and varied between the different types of substratum due to substratum choice. G. tigrinus did not show any behaviour indicative of interference competition in the presence of E. ischnus, and neither did E. ischnus or D. villosus in the presence of any of the other gammarideans. Swimming in the water layer may already enable E. ischnus to minimise its encounters with the stone-dwelling D. villosus and G. tigrinus. To maximise the encounters between E. ischnus and D. villosus, a fish (Lepomis gibbosus) was added to occupy the water layer during the aquarium experiments. E. ischnus showed a higher mortality in the presence of both D. villosus and fish, probably due to increased stress, as shelter opportunities to escape the predators had been minimised. The study shows that interference competition between gammaridean species can explain the replacement of the North American invader G. tigrinus by D. villosus in the river Rhine. E. ischnus and D. villosus both Ponto-Caspian invaders did not show interference competition in our experiments and co-exist in the Rhine.     

Relative abundance of Ponto‐Caspian gobiids in the lower Vistula River (Poland) 3‐ to 4 years after first appearance

In the years 2000–2001, specimens of two Ponto‐Caspian gobiids (racer goby Neogobius gymnotrachelus and monkey goby Apollonia fluviatilis) were recorded for the first time in the lower Vistula River, which is a part of the central inland corridor used by Ponto‐Caspian species to migrate in Europe. In 2004, seine net surveys were made in the river course to examine relative abundance of the non‐indigenous gobiids in the local small‐fish community. Altogether 4420 specimens with total lengths ranging from 10 to 340 mm (mostly <100 mm) belonging to 18 species were recorded in 70 hauls. Monkey goby was one of the subdominant species (18.1% of total number of fish captured; 64.3% frequency of occurrence in hauls) together with bleak Alburnus alburnus (21.0, 54.3), roach Rutilus rutilus (18.2, 67.1) and three‐spined stickleback Gasterosteus aculaetus (17.3, 37.1). Racer goby was less abundant but quite common (2.5, 32.9), similar to common bream/white bream Abramis brama/Blicca bjoerkna (6.4, 47.1), Eurasian perch Perca fluviatilis (5.7, 57.1), dace Leuciscus leuciscus (4.2, 40.0) and European bitterling Rhodeus sericeus amarus (3.4, 27.1). The study shows that Ponto‐Caspian gobiids dispersed successfully in the lower Vistula, becoming species common to its nearshore zone 3‐ to 4 years after their first appearance.     

Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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A new cardiid bivalve from the Pliocene Baklan Basin (Turkey) and the origin of modern Ponto‐Caspian taxa

Abstract: We present the first record of the cardiid genus Monodacna from the Pliocene of Anatolia, Turkey. Monodacna imrei sp. nov. was found in the Pliocene Killik Formation from the western margin of the Baklan Basin, in very marginal brackish to freshwater lacustrine deposits. The new record extends the stratigraphic range of the modern Ponto‐Caspian genus back into the Pliocene and adds to earlier evidence that modern Ponto‐Caspian taxa originated in the Pliocene of south‐western Turkey.     

Life History Traits of the Invader Dikerogammarus villosus (Crustacea: Amphipoda) in the Moselle River,France

The latest threatening invader in European freshwaters is Dikerogammarus villosus, a large gammarid of Ponto‐Caspian origin exhibiting a predatory behaviour. Its biology and population dynamics were studied over a one‐year period in a recipient ecosystem to determine bio/ecological traits having facilitated its rapid establishment. The study revealed that D. villosus reaches sexual maturity early, at six mm in length, and produces three reproductive peaks, though the species reproduces all year long, hence reflecting its multivoltine character. The study also revealed a female‐biased sex ratio, exceptional growth rates of up to 2.6 mm in two‐weeks in spring, and one of the highest fecundities of Western Europe gammarids. D. villosus exhibits a biological profile suggesting that only a few individuals can rapidly establish a new population in a recipient ecosystem, and allow this gammarid to become cosmopolitan in the near future. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Phylogeography and taxonomic status of trout and salmon from the Ponto‐Caspian drainages,with inferences on European Brown Trout evolution and taxonomy

Current taxonomy of western Eurasian trout leaves a number of questions open; it is not clear to what extent some species are distinct genetically and morphologically. The purpose of this paper was to explore phylogeography and species boundaries in freshwater and anadromous trout from the drainages of the Black and the Caspian Seas (Ponto‐Caspian). We studied morphology and mitochondrial phylogeny, combining samples from the western Caucasus within the potential range of five nominal species of trout that are thought to inhabit this region, and using the sequences available from GenBank. Our results suggest that the genetic diversity of trout in the Ponto‐Caspian region is best explained with the fragmentation of catchments. (1) All trout species from Ponto‐Caspian belong to the same mitochondrial clade, separated from the other trout since the Pleistocene; (2) the southeastern Black Sea area is the most likely place of diversification of this clade, which is closely related to the clades from Anatolia; (3) The species from the Black Sea and the Caspian Sea drainages are monophyletic; (4) except for the basal lineage of the Ponto‐Caspian clade, Salmo rizeensis, all the lineages produce anadromous forms; (5) genetic diversification within the Ponto‐Caspian clade is related to Pleistocene glacial waves; (6) the described morphological differences between the species are not fully diagnostic, and some earlier described differences depend on body size; the differences between freshwater and marine forms exceed those between the different lineages. We suggest a conservative taxonomic approach, using the names S. rizeensis and Salmo labrax for trout from the Black Sea basin and Salmo caspius and Salmo ciscaucasicus for the fish from the Caspian basin.     

Variation in fish community structure, richness, and diversity in 56 Danish lakes with contrasting depth, size, and trophic state: does the method matter?

Organisms inhabiting shallow near-shore waters are at risk of desiccation during water level fluctuations. Using laboratory experiments, we investigated the survival and behavioural defences of four freshwater amphipod species during substratum drying: three Ponto-Caspian invaders (Pontogammarus robustoides, Dikerogammarus haemobaphes and Dikerogammarus villosus) and the native Gammarus fossarum. We hypothesized that they would be able to survive air exposure events as well as to adjust their behaviour by following the decreasing water level and/or burying in the sediments. To test these hypotheses, we examined survival of each species on gradually drying sandy substratum as well as their horizontal and vertical migration behaviours. P. robustoides was most resistant to substratum drying and was the only species burying into the substratum. On the other hand, G. fossarum exhibited distinct horizontal migrations following the retreating waterline. These two species seem to be particularly well adapted to the drying environment. Defence mechanisms of D. haemobaphes and D. villosus were less efficient, though the former species also followed the retreating waterline to some extent. Our study demonstrates that exotic and native gammarids have several adaptations that enable them to invade and persist in habitats experiencing common water level fluctuations.     

Differential infection of exotic and native freshwater amphipods by a parasitic water mold in the St. Lawrence River

A parasitic water mold (oomycete) of unknown origin was discovered infecting live native and exotic freshwater gammarid amphipods (Gammarus fasciatus and Echinogammarus ischnus, respectively) in the upper St. Lawrence River. Infections were associated with rapid die-offs of natural populations of amphipods, especially the exotic E. ischnus. Analysis of sequences of three different segments of the rRNA gene cluster indicated that the parasite was in the Saprolegniaceae family, and is related to other crustacean-associated Saprolegniaceae. Specific primers were designed based on the SSU rRNA gene and utilized for semi-quantitative analysis of parasite presence in live and dead amphipods. In laboratory experiments, infection prevalence was higher in E. ischnus individuals than in native amphipods. In addition, dead E. ischnus individuals exhibited more intense infections than G. fasciatus individuals. In contrast to the Great Lakes where E. ischnus has replaced G. fasciatus at many locations, the native species remains abundant in the St. Lawrence River more than a decade after invasion by E. ischnus in the late 1990s. We hypothesize that the parasite is facilitating the co-existence of the two amphipod species by reducing the abundance of E. ischnus in environments in which it might otherwise be dominant.     

Seasonal variability in the grazing potential of the invasive amphipod Gammarus tigrinus and the native amphipod Gammarus salinus (Amphipoda: Crustacea) in the northern Baltic Sea

Mesograzers are known to reduce the biomass of their host plant and modify the structure of the whole macrophyte community in many ecosystems. Thus, the introduction of an efficient mesograzer may destabilize macrophyte community and also affect the native grazers. We estimated how large proportion of macrophyte production are consumed by the alien gammarid G. tigrinus and the native gammarid G. salinus in the species poor ecosystem of the northern Baltic Sea. We analysed whether G. tigrinus consumes different diet as the native G. salinus and whether the effect of G. tigrinus on the survival of the native G. salinus is macrophyte species specific. Grazing experiments showed that there was a clear difference in the grazing rates of gammarids among the studied macrophyte species in summer and autumn but not in spring. The grazing rates were significantly higher in the prevailing macrophyte Pilayella littoralis as compared to other macrophytes. The grazing was inversely related to the diurnal net photosynthetic values of macrophytes. The gammarid amphipods potentially removed only a minor part of plant primary production except for summer and autumn when grazing of a few perennial species exceeded macrophyte production. Macrophyte species and presence of G. salinus had no effect on the survival of G. tigrinus. The presence of G. tigrinus, however, reduced the survival of the native gammarids within P. littoralis in summer. To conclude it is likely that both native and alien gammarid amphipods do not exert significant pressure on the macroalgal communities in the northern Baltic Sea. Competitive interactions between G. tigrinus and G. salinus within the prevailing macrophyte P. littoralis is the likely explanation of the decline of the native gammarid amphipods after the establishment of G. tigrinus in the northern Baltic Sea.