Invader–invader interactions in relation to environmental heterogeneity leads to zonation of two invasive amphipods, Dikerogammarus villosus (Sowinsky) and Gammarus tigrinus Sexton: amphipod pilot species project (AMPIS) report 6

As biological invasions continue, interactions occur not only between invaders and natives, but increasingly new invaders come into contact with previous invaders. Whilst this can lead to species replacements, co-existence may occur, but we lack knowledge of processes driving such patterns. Since environmental heterogeneity can determine species richness and co-existence, the present study examines habitat use and its mediation of the predatory interaction between invasive aquatic amphipods, the Ponto-Caspian Dikerogammarus villosus and the N. American Gammarus tigrinus. In the Dutch Lake IJsselmeer, we found broad segregation of D. villosus and G. tigrinus by habitat type, the former predominating in the boulder zone and the latter in the soft sediment. However, the two species co-exist in the boulder zone, both on the short and longer terms. We used an experimental simulation of habitat heterogeneity and show that both species utilize crevices, different sized holes in a plastic grid, non-randomly. These amphipods appear to optimise the use of holes with respect to their ‘C-shape’ body size. When placed together, D. villosus adults preyed on G. tigrinus adults and juveniles, while G. tigrinus adults preyed on D. villosus juveniles. Juveniles were also predators and both species were cannibalistic. However, the impact on G. tigrinus of the superior intraguild predator, D. villosus, was significantly reduced where experimental grids were present as compared to absent. This mitigation of intraguild predation between the two species in complex habitats may explain the co-existence of these two invasive species.     

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.     

Interference competition among native and invader amphipods

Aquarium experiments were used to study indications of interference competition, such as substratum choice shifts, swimming activities and mortality of invasive and indigenous gammarids in each other's presence. The more recent invaders Gammarus tigrinus and Dikerogammarus villosus were more likely to prefer stone substratum, whereas the native Gammarus pulex and an earlier invader Gammarus roeseli were found more frequently in the water layer. Sand was the least likely substratum to be chosen by any of the species. G. pulex and G. roeseli did not alter their substratum preference in each other's presence. In the presence of D. villosus, G. pulex shifted towards smaller stones and increased its swimming activities, whereas D. villosus did not change its behaviour in the presence of G. pulex. These shifts may indicate interference competition, with D. villosus being the stronger competitor. The greatest shifts in substratum preference arose when one species had occupied a substratum before the other one was introduced, especially when D. villosus was already present before G. pulex was introduced, possibly indicating pre-emptive competition. Swimming activities of G. pulex increased in the presence of D. villosus, whereas D. villosus spent little time swimming. Mortality was comparable between the different experiments without any indication of predation. The effect of Intra Guild Predation (IGP) may not be reflected adequately by short-time experiments as moults occurred seldom during the experiments. Although no IGP was observed during our experiments, habitat shifts occurred, which may indicate that competitive interactions are apparent before IGP starts. Such shifts may serve to avoid intraguild competition.     

Native and exotic Amphipoda and other Peracarida in the River Meuse: new assemblages emerge from a fast changing fauna

Samples issued from intensive sampling in the Netherlands (1992–2001) and from extensive sampling carried out in the context of international campaigns (1998, 2000 and 2001) were revisited. Additional samples from artificial substrates (1992–2003) and other techniques (various periods) were analysed. The combined data provide a global and dynamic view on the Peracarida community of the River Meuse, with the focus on the Amphipoda. Among the recent exotic species found, Crangonyx pseudogracilisis regressing, Dikerogammarus haemobaphesis restricted to the Condroz course of the river, Gammarus tigrinusis restricted to the lowlands and seems to regress, Jaera istriis restricted to the ‘tidal’ Meuse, Chelicorophium curvispinumis still migrating upstream into the Lorraine course without any strong impact on the other amphipod species. After a rapid expansion Dikerogammarus villosushas continued its upstream invasion between 1998 and 2002 at a rate of 30–40 km per year, but no further progression was noticed in 2003. Locally and temporarily the native species (Gammarus fossarum and G. pulex) and naturalized species (G. roeseliand Echinogammarus berilloni)mayhave been excluded by the most recent invaders (mainly D. villosus), but none of the native and naturalized species has disappeared completely. Therefore, the number of amphipod species found in the River Meuse has increased. Moreover, the native and naturalized species keep on dominating the tributaries from which the recent invaders seem to be excluded. A changing Peracarida community structure is observed along the course of the River Meuse: four native or naturalized species inhabit the upstream (Lorraine) course, three invasive species dominate in the middle reach (Ardenne-Condroz zone), one exotic species is housed in the Border Meuse and three or four invasive species dominate the assemblages in the lowlands.     

Evaluation of the use of Water Framework Directive typology descriptors, reference sites and spatial scale in macroinvertebrate stream typology

The Rhine ecosystem is highly influenced by anthropogenic stresses from pollution, intensive shipping and increased connectivity with other large European rivers. Canalization of the Rhine resulted in a reduction of heterogeneity to two main biotopes: sandy streambeds and riverbanks consisting of groyne stones. Both biotopes are heavily subjected to biological invasions, affecting the rivers food web structure. The Ponto-Caspian amphipods, Chelicorophium curvispinum and Dikerogammarus villosus, have exerted the highest impact on this food web. The filterfeeding C. curvispinum dominated the Rhine food web on the stones in 1998, swamping the stone substrata with mud. However, in 2001 it decreased in numbers, most likely due to top-down regulation caused by increased parasitic and predatory pressure of other more recently invaded Ponto-Caspian species. D. villosus showed a fast population increase after its invasion and particularly influenced the macroinvertebrate community on the stones by predaceous omnivory. This species seemed to have maintained its predatory level after its population established. Effects of these mass invaders on the macroinvertebrate community of sandy streambeds in the Rhine are unclear. Here, low densities of macroinvertebrates were observed with the Asiatic clam, Corbicula fluminea, as most abundant species. Stable isotope values of food webs from the stones and sand in 2001 were similar. Aquatic macrophytes are nearly absent and the food web is fuelled by phytoplankton and particulate organic matter, originating from riparian vegetation as indicated by similar δ¹³C values. Omnivores, filter-, deposit-, and detritus-feeders are the primary and secondary macroinvertebrate consumers and function as keystone species in transferring energy to higher trophic levels. Invaders comprise 90% of the macroinvertebrate numbers, and can be considered ecosystem engineers determining the functional diversity and food web structure of the Rhine by either bottom-up or top-down regulation.     

Feeding rates,assimilation efficiencies and growth of two amphipod species on biodeposited material from zebra mussels

1. Accumulation of organic material by the zebra mussel Dreissena polymorpha is assumed to be the source of a biodeposition‐based food web. However, only little is known about the importance of the biodeposited material as a food source and its contribution to increased abundances of macroinvertebrates in the presence of D. polymorpha. 2. Feeding, assimilation and growth of the amphipods Gammarus roeselii and Dikerogammarus villosus on food sources directly and indirectly associated with D. polymorpha (biodeposited material and chironomids) and on conditioned alder leaves were measured. The stoichiometry of carbon, nitrogen and phosphorus of the diets was measured as an important determining factor of food quality. 3. Chironomids had the highest nitrogen and phosphorus contents, alder leaves were depleted in nitrogen and phosphorus, and the stoichiometry of biodeposited material was intermediate. 4. Both amphipod species had highest feeding rates and assimilation efficiencies on chironomids. Gammarus roeselii fed more on biodeposited material than on alder leaves, but assimilation efficiencies were similar; D. villosus also had similar feeding rates and assimilation efficiencies on the two diets. 5. Both amphipod species had highest growth rates on chironomids and lowest growth rates on alder leaves. Both grew at intermediate rates on biodeposited material of D. polymorpha. The growth rates of the amphipod species were related to food stoichiometry. Overall, the invasive D. villosus grew faster than the indigenous G. roeselii. 6. Food resources directly and indirectly associated with D. polymorpha are potential diets for amphipods, providing further evidence for a D. polymorpha biodeposition‐based food web.     

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.     

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.     

Planktonic bloom of an introduced species of Polysiphonia (Ceramiales,Rhodophyta) along the coast of North Carolina,USA

Analysis was made of the species composition, egg number per female and the size-frequency of Amphipoda living on the dominating submerged macrophytes (Potamogeton perfoliatus and Myriophyllum spicatum) at 10 sampling stations of the northern and southern shoreline of Lake Balaton. The dominating Amphipoda at each sampling station was Corophium curvispinum (85.9–99.8%, mean: 96.6%). Besides this, two other species — Dikerogammarus haemobaphes and D. villosus — were also found in the samples. The two Dikerogammarus species were found in highest percentage near Keszthely (14.1 %); at the other places studied they were under 5%. The number of Amphipoda individuals per g macrophyte dry weight ranged from 5 to 574; the lowest value was found near Keszthely, the highest near B. Mária. The developmental stages of the C. curvispinum population differ in different parts of Lake Balaton: the adult (male and female) specimens occurred in highest percentage near Keszthely. The mean number of eggs per C. curvispinum female ranged from 2.4 to 6.3, showing differences at the different sampling stations. Regarding the two Dikerogammarus species, there were many more D. haemobaphes than D. villosus individuals at almost every station (mean: 75%). The mean number of eggs per egg-carrying female of D. haemobaphes was 11.7, being 19.1 for D. villosus.The total biomass of amphipods (in mg animal dry weight/g macrophyte dry weight) ranged from 1.2 to 59.8. The lowest value was observed near Keszthely (the most hypertrophic basin of Lake Balaton), the highest value near B. Maria     

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.     

Feeding choice and predation pressure of two invasive gammarids, <Emphasis Type="Italic">Gammarus tigrinus</Emphasis> and <Emphasis Type="Italic">Dikerogammarus villosus</Emphasis>, under increasing temperature

In most European freshwater ecosystems, the invasive gammarids Gammarus tigrinus and Dikerogammarus villosus strongly impair recipient communities through predation of a wide range of native invertebrates. Due to the effects of temperature on both the physiology and the behaviour of such ectotherms, understanding how global warming may influence their ecological impact is a research priority. These species were therefore exposed to three different food types to determine their detritivorous, herbivorous and carnivorous characters, and predation was measured characterizing the Holling’s functional response. The effect of increasing water temperatures (15, 20, 25°C) on both the food choice and predatory activities was investigated. Both species showed a significant preference for animal tissues at all temperatures. The total food intake increased with temperature for G. tigrinus but did not change for D. villosus, which may result from specific species differences in metabolic requirements. The consumption of live prey strongly increased with temperature. The main differences were an increased searching efficiency in G. tigrinus and a decreased handling time in D. villosus as temperature increased, which may result from differences in foraging strategies. These results suggest that climate change is likely to increase the predation pressure of both invasive gammarids on prey species.     

Biodiversity and Structure of Macroinvertebrate Communities Along a Small Permanent Salinity Gradient (Meurthe River,France)

Changes in the macroinvertebrate community were investigated over 10 months at four sites along a 19 km salinity gradient (0.21–2.60 g l⁻¹) in a sixth-order stream, the Meurthe River, northeastern France. Abiotic characteristics other than salinity were similar between the sites. Macroinvertebrate taxonomic richness decreased by 30% downstream of the 1.4 g l⁻¹ sites while diversity, evenness or total abundance of taxa did not change along the gradient. In terms of functioning, a slight change in relative abundances of invertebrate feeding groups followed the salinity gradient. Eight invertebrate assemblages occurred within specific salinity distributions were identified. The exotics Gammarus tigrinus, Dreissena polymorpha, Corbicula fluminalis and Corophium curvispinum, were more abundant at the highest salinity site. These results suggest that rising salinity concentrations drastically affect the species composition, including favouring exotic species.     

Mutual predation between and cannibalism within several freshwater gammarids: Dikerogammarus villosus versus one native and three invasives

Dikerogammarus villosus, a Ponto-Caspian species, is known to be a particularly successful invader, and is currently the prevailing invasive gammarid in Southern German large waters. Field observations suggest that D. villosus has replaced the native G. pulex and the invasive D. haemobaphes, also a Ponto-Caspian species, in some reaches of the German Danube. Dikerogammarus villosus is also believed to be the reason why Echinogammarus berilloni, a Mediterranean species, and Dikerogammarus bispinosus, a further Ponto-Caspian gammaridean species, could not build significant populations in Southern German rivers. Because intraguild predation (IGP) is regarded as a major force leading to species displacements, we hypothesized that superiority in predation by D. villosus is responsible for the disappearance or limited occurrence of several native and invasive gammarideans in many reaches of Southern German waters. To test this hypothesis, we conducted a series of laboratory IGP experiments with 1:1 combinations in which D. villosus was allowed to prey on other gammarids and vice versa. We also studied the extent of cannibalism within species. Dikerogammarus villosus was a stronger predator than G. pulex and E. berilloni. However, D. villosus was not stronger in predation than its relatives D. haemobaphes and D. bispinosus, although results with the combinations of D. villosus and D. bispinosus were less conclusive, especially in the female–female combination. Cannibalism rate was high in D. haemobaphes compared with other species. We conclude that superiority in IGP experiments could explain the disappearance of G. pulex and the missing or limited occurrence of E. berilloni since the arrival of D. villosus in Southern German rivers. However, the results of IGP experiments do not explain why D. haemobaphes was displaced by D. villosus in many places and why D. bispinosus could not build significant populations in Southern German waters. Possibly specific combinations between mutual predation and cannibalism or factors other than predation could have affected displacement and occurrence patterns of Dikerogammarus species in Southern German waters.     

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.     

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.     

The survival and growth of Gammarus tigrinus Sexton (Crustacea: Amphipoda) in relation to salinity and temperature

Field collections from an inland saline lake indicated that a rapid increase in numbers of a population of Gammarus tigrinus was positively correlated with high summer temperatures and salinities. The laboratory experiments on growth are an attempt to determine whether high summer temperatures or optimum salinity were primarily responsible for the rapid increase in numbers. Furthermore, G. tigrinus was not found in either of two adjacent lakes; one non-saline, the other at a higher salinity. The laboratory experiments on survival are an attempt to determine whether the restricted distribution of G. tigrinus was related to salinity. The implications of the field and laboratory studies are discussed and compared with other data from Europe and North America.     

Potential effects of the invasive ‘killer shrimp’ (Dikerogammarus villosus) on macroinvertebrate assemblages and biomonitoring indices

1. Water quality monitoring data from 10 watercourses and laboratory mesocosm studies were used to assess the potential impacts of the crustacean amphipod invader Dikerogammarus villosus on resident macroinvertebrate assemblage structure in Central European fresh waters. 2. The presence of D. villosus was associated with a decline in the prevalence of many native species, pollution sensitive as well as pollution tolerant, and changes in biotic indices, despite the trends of improved water quality coinciding with the invasion period. A general increase in the prevalence of other invaders was also noted. The potential impacts of D. villosus were substratum dependent, differing between stone, concrete and sand‐dominated sites. 3. Mean Multimetric Macroinvertebrate Index Flanders (MMIF) values were marginally lower when D. villosus was present (P < 0.06), as opposed to when other amphipod species or no amphipods were present, despite the improved water quality. Mesocosm studies showed that several macroinvertebrate taxa were completely eliminated in treatments with D. villosus, oligochaete worms, Caenidae mayfly, chironomids and tipulids being particularly vulnerable to D. villosus predation. Biological Monitoring Working Party (BMWP) scores were lower in mesocosms with D. villosus as opposed to the native Gammarus pulex or no amphipods at all. 4. We predict that resident macroinvertebrate assemblages in both Central Europe and Britain will come under increasing pressure as D. villosus invasions progress. Consequently, macroinvertebrate biotic indices, such as the MMIF or BMWP, may need to be revised to account for changes in taxa sensitivities to water quality as well as increased predation and competition.     

Influence of substrate preference and complexity on co-existence of two non-native gammarideans (Crustacea: Amphipoda)

Substrate choice, swimming activity and risk to predation by burbot (Lota lota) of the well established Gammarus roeselii and the invader Dikerogammarus villosus were studied in mixed and single-species aquarium experiments. We used stones, gravel and aquatic weeds (Elodea, Chara) as substrates. We hypothesized that both species have different substrate preferences and that substrate affects the predation risk. We also assumed that presence of D. villosus influences substrate preference and predation risk of G. roeselii since the invader is known to affect the behavior of other gammarids. Adults of D. villosus in single species experiments and juveniles in mixed and single species experiments were evenly distributed over the different substrates but adults in mixed species experiments were more likely to prefer stone substrate. In contrast, adults and juveniles of G. roeselii clearly preferred aquatic weeds independent of the presence/absence of the invader. Both species preferred substrates with fissured surface over substrates with smooth surface. Gammarus roeselii was observed swimming more often than D. villosus in the open water but its swimming activity was lower when its preferred substrate was present compared with its swimming activity if non-preferred substrates were present. Predation rate of burbot on D. villosus was comparatively low and independent of the substrate. Burbot consumed many more G. roeselii than D. villosus, both in mixed and single species experiments. But when the preferred substrate of G. roeselii (weeds) was used in the experiments, predation rate of burbot on G. roeselii was somewhat lower than that when non-preferred substrates were present. The results of the experiments support our hypothesis that the gammarids studied here have different substrate preferences and that presence of the preferred substrate can affect predation risk. However, there is no evidence that presence of D. villosus affected substrate choice or predation risk in G. roeselii. We consider that differences in use of spatial niches permit co-existence of G. roeselii and D. villosus in the wild when substrates are diverse. The fact that G. roeselii than D. villosus is more often observed swimming in the open water may explain its higher risk of being captured by fish.     

Distribution and aspects of population dynamics of the zebra mussel,Dreissena polymorpha (Pallas, 1771), in the lake IJsselmeer area (The Netherlands)

Summary The distribution, settling and growth of the zebra mussel, Dreissena polymorpha Pallas, in Lake IJsselmeer and Lake Markermeer in The Netherlands were studied from 1980 to 1985. In these lakes D. polymorpha is the most important food source for wintering diving ducks. The study was part of an investigation into the carrying capacity of the lakes for these birds.     

The population ecology of gammarus tigrinus (sexton) in the reed beds of the Tjeukemeer

The population ecology of Gammarus tigrinus (Sexton) was studied in the Tjeukemeer during 1969 and 1970. G. tigrinus reaches very high densities — up to 24,000/m² in parts of the study area. In 1970, the summer densities were 2–21/2 times greater than in 1969. Individuals do not grow to such large sizes in the summer as at other times of the year. Females begin to carry eggs in March or April and reproduction ceases in November. Large females have larger broods than smaller animals and the average size of the brood varies with the time of year. The egg incubation period and growth rate are dependent upon temperature. At summer temperatures females became sexually mature after about four weeks and the egg incubation period is about io days. The entire population is turned over about three times during the year. A combination of rapid growth rate, early onset of sexual maturity and high fecundity are probably responsible for the rapid spread of G. tigrinus throughout much of the Netherlands.