Differences in composition of macroinvertebrate communities with invasive and native Gammarus spp. (Crustacea: Amphipoda)

SUMMARY 1. Assessing the effects on communities of invasive species is often confounded by environmental factors. In Irish rivers, the introduced amphipod Gammarus pulex replaces the native G. duebeni celticus in lowland stretches. The two amphipods are associated with different macroinvertebrate communities, which may in part be the result of natural longitudinal physicochemical change. However, this hinders assessment of any direct community impacts of the invasive as compared with the native species. Here, we report on a fortuitous circumstance that allowed us to uncouple the community effects of Gammarus species from environmental differences.
2. The lowland stretch of the River Lissan is dissected by a weir, which has slowed the upstream invasion by G. pulex . We took physicochemical measurements and macroinvertebrate samples from three contiguous 150 m reaches of this stretch with G. pulex only, mixed Gammarus and G. d. celticus only communities.
3. We found no biologically significant differences in physicochemistry among the three reaches. Overall invertebrate densities did not differ among reaches. However, G. pulex numerically dominated its reach, whilst G. d. celticus abundance was relatively low in its reach. The G. pulex reach had significantly higher overall biomass because of high invader abundance. In addition, both diversity and species richness of macroinvertebrate communities were lower in the G. pulex than the G. d. celticus reach, with the mixed Gammarus reach intermediate.
4. Ordination indicated distinctly different associations of invertebrate community samples and taxa that were best explained by the distributions of the Gammarus species. Again, the mixed Gammarus species samples were intermediate.
5. This study indicates that the invasive G. pulex has a greater impact on macroinvertebrate community composition than the native G. d. celticus , probably through biotic interactions such as predation.     

Differential microdistributions and interspecific interactions in coexisting Gammarus and Crangonyx amphipods

In freshwaters in Northern Ireland, several amphipods occur, with the native Gammarus dubeni celticus and introduced Gammarus pulex and Crangonyx pseudogracilis often being found together in the same river and lake systems. We examined what may happen when the three amphipod species co-occur within the same patch of lake or pooled area of a river. We conducted a laboratory simulation of a lake/river habitat, and this showed that G. d. celticus and G. pulex exhibited very similar distribution patterns to one another, when presented with the same complex habitat template. Such microdistribution patterns contrasted markedly with the microhabitat usage by C. pseudogracilis. Cannibalism was low for all species in both single and mixed species treatments. Intraguild predation (IGP) was also very low on both G. pulex and G. d. celticus when all three species co-occurred. However, C. pseudogracilis suffered heavy IGP from both Gammarus species, with 33% of the tank population of C. pseudogracilis being eliminated in the presence of Gammarus within only 12 h. Such IGP may account for C. pseudogracilis occurring more frequently and in greater abundance in patches of natural lotic and lentic systems where predatory Gammarus are either absent or scarce.     

Introduction of the non-indigenous amphipod Gammarus pulex alters population dynamics and diet of juvenile trout Salmo trutta

Summary 1. The amphipod Gammarus pulex , introduced to Irish rivers with the aim of enhancing trout feeding, is displacing the native Gammarus duebeni celticus . These two species are generally associated with different environmental conditions and macroinvertebrate communities, confounding assessment of effects of the invader as compared with the native on fish populations. Here, we uncouple effects of the two Gammarus species from environmental gradients.
2. A weir dissects a lowland stretch of the River Lissan, slowing the upstream invasion by G. pulex and resulting in contiguous G. pulex , mixed species and G. d. celticus reaches. Total invertebrate abundance and biomass in the benthos were significantly higher in the G. pulex reach, driven by high invader abundance, with low abundance of other taxa. Gammarus pulex was particularly prominent in night-time drift.
3. Correspondingly, densities and biomass of 0+ trout were significantly higher in the G. pulex reach, while instantaneous loss rates were lower. Fish growth rates were similar among the three reaches.
4. In the G. pulex reach, this invader dominated the diet of 0+ trout, leading to ingestion of significantly higher invertebrate biomass than fish in the other reaches. Fish generally preyed on Gammarus in proportion to its abundance, but exhibited some positive selection for G. pulex in the invaded reach.
5. The negative effects of the invader on native invertebrates are contrasted with positive effects on juvenile trout. This indicates changes in energy flux after invasion, with differential resource use or assimilation by G. pulex probable underlying mechanisms. As the frequency of amphipod invasions increases globally, investigations of their role as strong interactors at multiple levels of ecological organisation is required if the consequences of deliberate and unintentional introductions are to be predicted, and ultimately, prevented.     

Predation on mayfly nymph, Baetis rhodani, by native and introduced Gammarus: direct effects and the facilitation of predation by salmonids

1. In a series of laboratory experiments, we assessed the predatory nature of the native Irish amphipod, Gammarus duebeni celticus , and the introduced G. pulex , towards the mayfly nymph Baetis rhodani . We also investigated alterations in microhabitat use and drift behaviour of B . rhodani in the presence of Gammarus , and indirect predatory interactions with juvenile Atlantic salmon, Salmo salar .
2. In trials with single predators and prey, B. rhodani survival was significantly lower when Gammarus were free to interact with nymphs as than when Gammarus were isolated from them. The invader G. pulex reduced the survival of B. rhodani more rapidly than did the native G. d. celticus . Both Gammarus spp. were active predators.
3. In `patch' experiments, B. rhodani survival was significantly lower both when G. pulex and G. d. celticus were present, although the effect of the two Gammarus species did not differ. Again, active predation of nymphs by Gammarus was observed. Significantly more nymphs occurred on the top and sides of a tile, and per capita drifts were significantly higher, when Gammarus were present. Baetis rhodani per capita drift was also significantly higher in the presence of the introduced G. pulex than with the native G. d. celticus .
4.  Gammarus facilitated predation by salmon parr of B. rhodani by significantly increasing fish–nymph encounters on exposed gravel and in the drift. There were no differential effects of the two Gammarus spp. on fish – B . rhodani encounters or consumption.
5. We conclude that Gammarus as a predator can have lethal, nonlethal, direct and indirect effects in freshwaters. We stress the need for recognition of this predatory role when assigning Gammarus spp. to a `Functional Feeding Group'.     

The nature and implications of differential predation between Gammarus pulex and G. duebeni celticus (Crustacea: Amphipoda)

The mutual predatory behaviour between Gammarus pulex (Linnaeus, 1758) and Gammarus duebeni celticus (Stock & Pinkster, 1970) is described. The implications of the predatory nature of these species are discussed in terms of the feeding ecology of Gammarus and other amphipods. Predation rates by males on moulted congeneric females are assessed and compared. When males are present at the moult of a congeneric female, 100%of females are eaten in both species/sex combinations. When females are allowed to recover from moulting prior to presentation to a congeneric male, predation rates are significantly reduced in both species. However, significantly more G. d. celticus females are eaten by G. pulex males compared with the reciprocal situation. The predatory 'clumping' behaviour of both species is described and shown to be a form of feeding frenzy upon congenerics. Clumping behaviour also results in significantly higher rates of congeneric predation on G. d. celticus females than on G. pulex females. Support is thus given to the hypothesis that differential predation by males on moulted females may be the primary mechanism by which the introduced G. pulex has displaced the native Irish species G. d. celticus . The implications of amphipod congeneric predation are discussed in terms of observed patterns of interspecific coexistence and exclusion.     

Invasion by the amphipod Gammarus pulex alters community composition of native freshwater macroinvertebrates

Assessing the effects of invading species on native community structure is often confounded by environmental factors and weakened by lack of replicated, long-term pre- and post-invasion monitoring. Here, we uncouple the community effects of a freshwater amphipod invader from environmental differences. In Irish rivers, the introduced Gammarus pulex replaces the native Gammarus duebeni celticus . However, the River Lissan in Northern Ireland is dissected by a weir that has slowed the upstream invasion by G. pulex . This allowed us in 2000 to sample three contiguous 150-m reaches that were (1) G. pulex dominated; (2) mixed Gammarus spp.; and (3) G. duebeni celticus only. In 2003, we resampled these reaches and one additional of mixed Gammarus species and one with only G. duebeni celticus further upstream. In temperature, conductivity, and pH, there were statistically significant but no biologically relevant differences among the five reaches of 2003, and between the three reaches surveyed in both years. Although there was evidence of recovery in macroinvertebrate diversity and richness in invaded reaches between years, continued upstream invasion was associated with sustained reductions in these community metrics as compared to un-invaded sites. Community ordination indicated (1) different associations of community composition attributed to the distribution, abundance, and biomass of the invader; and (2) increasing similarity of invaded communities over time. The impact mechanisms of G. pulex on macroinvertebrate community composition may include predation and competition. The consequences of the observed community changes for ecosystem functioning require further investigation.     

Parasitic infection manipulates sodium regulation in the freshwater amphipod Gammarus pulex (L.)

The acanthocephalan parasite Polymorphus minutus induces both physiological and behavioural effects in its intermediate host, Gammarus pulex. The net effect of parasite infection is to increase the likelihood of transmission to the definitive host. Osmoregulation is an energetically expensive mechanism that allows G. pulex to survive in dilute media. Any factor influencing osmoregulation is thus likely to affect the allocation of resources to other areas. This study investigated whether P. minutus infection alters sodium regulation in G. pulex. Haemolymph sodium concentration, water permeability and sodium fluxes were measured over the salinity acclimation range of G. pulex. Water permeability was unaltered by either acclimation salinity or parasite infection. Acclimation to 12‰ significantly raised the haemolymph sodium concentration, reduced the sodium influx, and increased the sodium efflux, to the same extent in both uninfected and infected G. pulex. However, parasite infection induced a significant increase in haemolymph sodium concentration in G. pulex acclimated to 6‰, which was not observed in uninfected G. pulex acclimated to the same salinity. Also, both sodium influx and sodium efflux were significantly lower in parasitized G. pulex acclimated to 6‰, when compared to uninfected G. pulex acclimated to the same salinity. It was concluded that the parasite induced disturbances to sodium regulation in G. pulex acclimated to 6‰ were a functional consequence of the manipulative strategy employed to alter behaviour, rather than a primary target.     

Pairing success and sperm reserve of male Gammarus pulex infected by Cyathocephalus truncatus (Cestoda: Spathebothriidea)

Manipulative parasites with complex life cycles are known to induce behavioural and physiological changes in their intermediate hosts. Cyathocephalus truncatus is a manipulative parasite which infects Gammarus pulex as intermediate host. G. pulex males display pre-copulatory mate guarding as a response to male-male competition for access to receptive females. In this paper, we tested the influence that C. truncatus-infection might have on male G. pulex sperm number and pairing success. We considered 3 classes of G. pulex males in our experiments: (i) uninfected males found paired in the field, (ii) uninfected males found unpaired in the field, or (iii) infected males found unpaired in the field. Both infected males and uninfected unpaired males paired less with a new female than uninfected paired males did. Furthermore, infected males appear to be at a strong disadvantage when directly competing for females with a healthy rival male, and had fewer sperm in their testes. We discuss the potential effect of male and female mating strategies on such male host mating alteration.     

A comparison of numerical taxonomic techniques using measurements on the genera Gammarus and Marinogammarus (Amphipoda)

Numerical taxonomy involves the use of quantitative techniques to assess the similarity between species. In this work a comparative study has been made on seven of these methods. The data consisted of measurements made on thirteen amphipod species belonging to the genera Gammarus and Marinogammarus. A principal coordinate analysis, a canonical variate analysis and several methods of cluster analysis were used. There is also a discussion on the Uniquely derived character concept.
The main conclusion was that the numerical techniques give results which agree generally but certainly differ in detail. The consequences are discussed and certain methods are recommended. On the basis of the seven methods of analysis, a number of well-defined groups were recognized within the animals in the study. The genus Gammarus is reckoned to be distinct from the genus Marinogammarus. G. pulex and G. lacustris are similar to each other but rather different from the other Gammarus species. All the thirteen species are clearly differentiated although the two forms of G. duebeni are quite similar.     

Effects of temperature on persistence times of native and invasive gammarid species in the stomachs of Cottus gobio

Gastric lavage was used to investigate the effects of temperature on persistence time of two amphipod species, one native Gammarus pulex and one invasive Gammarus roeseli , in the stomachs of bullhead Cottus gobio . Persistence time was strongly influenced by temperature and prey type, such that G. pulex species degraded faster than G. roeseli .     

Lethal and sub-lethal effects of acid, aluminium and lime on Gammarus pulex during repeated simulated episodes in a Welsh stream

SUMMARY. 1. A soft-water stream in upland mid-Wales was dosed with sulphuric acid, aluminium sulphate and limestone slurry to produce a zone of aluminium at low pH. and one of aluminium, low pH and added lime.
2. Three episodes of 24-h duration were induced, each separated by a 12-h inter-dosing period, and the responses of the crustacean Gammarus pulex monitored throughout.
3. G. pulex parasitized by the acanthocephalan Pomphorhynchus laevis were significantly more sensitive than unparasitized individuals.
4. Liming significantly reduced mortality during dosing but postexposure deaths were greatest for animals which had been exposed in the time zone.
5. The disruption of precopulalory behaviour in G. pulex was rapid and liming did not mitigate this response.     

Factors affecting the microdistribution of Gammarus pulex (Amphipoda): an experimental study

SUMMARY. 1. The tnierodistribution of Gammarus species is size assortative: larger animals are associated with larger substrate particles. Using an artificial stream with scrubbed substrate we investigate the hypothesis that the assortative mechanism is stimulated by current avoidance rather than food searching.
2. In uniformly mixed substrate, G. pulex tend to move upstream, but in patchy substrate they are found in a predictable substrate grade. There is no discrimination between patches by different size classes of G. pulex in static water. The size assortative field pattern can only be reproduced under flow conditions in the laboratory. It occurs in the absence of food and irrespective of the sequence or location of substrate patches.
3. As density increases, G. pulex are displaced from the optimal patches but move into the open stream rather than into the cover of alternative patch types.
4. We discuss the significance of these results and suggest that some earlier models explaining mechanisms for microdistribution patterns tnay have been unnecessarily complex. Contrasts between natural and simulated stream situations and their experimental advantages are noted.     

The effect of the acanthocephalan parasite (Pomphorhynchus laevis) on the drift of its intermediate host (Gammarus pulex)

SUMMARY 1. The drift of Gammarus pulex in a population in which approximately 20% of adults are infected with the acanthocephalan parasite Pomphorhynchus laevis was monitored at margin and mid-river sites in the River Teme, England over a 24-h period in mid-summer.
2. Drift densities showed a diurnal pattern with a large increase at night, independent of parasite burden.
3. There was no significant effect of site (margin or mid-river) on the proportion of parasitized and unparasitized G. pulex found in the drift or the benthos.
4. The drift of parasitized G. pulex was significantly greater than unparasitized animals. Gammarids harbouring only one parasite were found in significantly higher proportions in the drift than those with two or more parasites.
5. At both sites (margin and mid-river) the proportion of unparasitized adult G. pulex in the drift was significantly lower than that in the benthos. However, there was a significantly higher proportion of parasitized animals in the drift than in the benthos.     

Carotenoids of two freshwater amphipod species (Gammarus pulex and G. roeseli) and their common acanthocephalan parasite Polymorphus minutus

Carotenoid compositions of two freshwater Gammarus species (Crustacea: Amphipoda) and of their common acanthocephalan parasite Polymorphus minutus were characterized. The effect of carotenoid uptake by the parasite was addressed by comparing the carotenoid content of uninfected and infected female hosts. Using high-pressure liquid chromatography (HPLC), co-chromatography of reference pigments and electron ionization mass spectrometry of collected HPLC fractions (EI-MS), several xanthophylls and non-polar compounds were identified. Seven kinds of carotenoids, mainly xanthophylls, were identified in gammarids. Astaxanthin was predominant, amounting to 40 wt.% of total carotenoid in both uninfected G. pulex and G. roeseli. By contrast, we found only non-polar compounds with a predominance of esterified forms of astaxanthin in P. minutus larvae. No significant effect of infection on carotenoid content was evidenced in G. pulex and G. roeseli females. Our study highlights the use of a Matrix Solid Phase Dispersion as an efficient extraction method of both xanthophylls and non-polar pigments in small samples, including lipid-rich ones as P. minutus parasite. We discuss on the presumptive pathway leading to the formation of free astaxanthin in gammarids via hydroxy compounds, and on the accumulation of esters of astaxanthin in parasites.     

Introduction experiments with Gammarus spp. (Crustacea: Amphipoda) in the Isle of Man (British Isles), 1949–1995

This study uses data from field introduction experiments with Gammarus spp. conducted by H. B. N. Hynes in the Isle of Man in the period 1949-1969, together with data from re-sampling programmes by the present authors in 1974/5, 1988 and 1995, to examine the interaction between the native freshwater amphipod G. duebeni and the introduced G. pulex. Although Hynes' deliberate introductions of G. pulex into streams in the Isle of Man at first seemed to give equivocal results, their examination in the light of subsequent evidence indicates that G. duebeni is, under some as yet unknown condition(s), sometimes able to withstand the impact of invasions by its competitive rival G. pulex. The presentation of Hynes' data is thus a belated but valuable contribution to knowledge in this field, although we must stress that such experiments involving transplantations of species should not be carried out owing to the ecological damage that can follow. In addition, we discuss our discovery of a truly exotic amphipod in the Isle of Man, the N. American Crangonyx pseudogracilis .     

Sexual dimorphism in Gammarus pulex: the effect of current flow on pre-copula pair formation

SUMMARY. 1. Male Gammarus pulex (L.) are typically twice as heavy as females in field caught pre-copula pairs and mating is positively size assortative. Earlier work had shown that relatively large males can swim in pre-copula at higher current speeds than males that are relatively small compared with their mates. We describe experiments on pair formation in the laboratory, in static water and in an artificial stream.
2. Pairs formed in the artificial stream had a significantly higher weight ratio than either those formed in static water or those caught in the field. In both experimental conditions, males in pairs were significantly larger than singles. In the stream, the weight ratio of pairs increased with formation sequence due to a significant decrease in the size of selected females.
3. It is concluded that these results are consistent with the hypothesis that the relatively large size of paired male G. pulex is a product of mechanical constraints rather than male–male competition.     

Physiological and behavioural responses of Gammarus pulex exposed to acid stress

Physiological and behavioural responses of the acid-sensitive amphipod Gammarus pulex exposed to a wide range of acid conditions (pH 4.1, 5.1, 6.0) under laboratory conditions were investigated. An exposure of 38 h to acid conditions caused significant decreases in survival rate, osmolality, haemolymph Na+ concentration, ventilatory and locomotor activity compared to organisms exposed to a circumneutral medium (pH 7.9). We highlighted the interest of using individual response distribution, since drastic effects can be detected in organisms exposed to pH 6.0, in particular for osmolality: The response can be divided into two groups, unimpacted and impacted organisms. Moreover this representation permitted to evaluate the health level of individual organisms through the determination of threshold values. Significant correlations between mean pH and mean physiological/behavioural responses were observed. The relationships between individual responses permitted not only to compare endpoints, but also to show that affected organisms were impacted by ionoregulation failure, hypoventilation and low locomotor activity. The energetic reallocation in favour of maintenance functions, such as osmoregulation, is discussed. The results of this study indicate that the values of haemolymph Na+ concentration, osmolality and locomotor activity in G. pulex could be effective ecophysiological/behavioural markers to monitor freshwater ecosystems and to assess the health of organisms or populations.     

The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)

This study examines the cannibalistic behaviour of the freshwater amphipods Gammarus duebeni celticus Stock & Pinkster, 1970 and G. pulex (L., 1758). In the first experiment, interactions were staged among all combinations of single adult males, single adult females, adults in the precopulatory mate-guarding phase and juveniles. Cannibalism by inter-moult individuals on newly moulted conspecifics occurred in all interaction categories in both species. Gammarus d. celticus , however, were significantly more cannibalistic than G. pulex. Cannibalism between and within sex and size categories (males > females > juveniles) was facilitated by the vulnerability of individuals at moult. Individuals of smaller size categories, however, did not cannibalize newly moulted conspecifics of larger size categories. Males were less cannibalistic on newly moulted females than on newly moulted males and juveniles and, when in the precopulatory condition, appeared to defend females from cannibalistic attacks. In a second experiment, stream conditions were simulated in the laboratory and replicated populations monitored for nine weeks. High levels of cannibalism, and the species and sex differences in cannibalism identified in the first experiment, were confirmed under these heterogeneous conditions. Cannibalism by males on their newly moulted female mating partners, termed 'reversed' sexual cannibalism, was further investigated. When males were deprived of foraging opportunities, cannibalism of precopulatory partners was significantly more frequent. The occurrence of 'reversed' sexual cannibalism is thus interpreted as a conflict between motivation to feed and motivation to mate.     

Influence of predator and dietary chemical cues on the behaviour and shredding efficiency of Gammarus pulex

In two laboratory experiments, we examined short- and long-term responses of the detritivorous amphipod Gammarus pulex to chemical cues from potential predators fed various diets. In the first experiment we studied the short-term effect on G. pulex (locomotory activity) when exposed to chemical cues from three co-existing predators; sculpin (Cottus gobio), trout (Salmo trutta), and signal crayfish (Pacifastacus leniusculus). Chemicals from sculpins and trout induced a short-term decrease in locomotory activity in G. pulex, whereas crayfish did not. There was no difference in activity between G. pulex exposed to water scented by trout or sculpin, and these responses were independent of predator diet (G. pulex, Asellus aquaticus and starved). In the second experiment we examined whether longer-term exposure (4 week) to chemical cues from sculpins affects rates of leaf processing by G. pulex. During the first week, G. pulex consumed significantly more leaves in the control (i.e., no fish cue) than in the fish cue treatment. After 4 weeks, however, there was no difference in total leaf processing rate between treatments indicating an adaptation to the cue.     

Distribution and microhabitats of native and non-native gammarids (Amphipoda, Crustacea) in Brittany, with particular reference to the endangered endemic sub-species Gammarus duebeni celticus

Aim  To assess temporal changes in gammarid distribution in Brittany and microhabitat-use overlap between the endangered endemic Gammarus duebeni celticus Stock & Pinkster, 1970 , the expanding natives G. pulex (Linnaeus, 1758) and Echinogammarus berilloni (Catta, 1878), and the introduced G. tigrinus Sexton, 1939.
Location  Brittany and adjacent regions in western France.
Methods  The spatial and temporal patterns in distribution of gammarids at the scale of Brittany were studied using 351 sites. Longitudinal distributions (from the source to the estuary of the river) and microhabitat-use (substratum type and water velocity) were also considered in selected rivers.
Results  At the regional scale, all species occurred together less often than expected statistically, with significant deviations from expected for G. pulex vs. both G. duebeni celticus and G. tigrinus , and for E. berilloni vs. both G. duebeni celticus and G. tigrinus . However, at the microhabitat scale, E. berilloni occurred significantly more often than expected with the endemic G. duebeni celticus , and this appears to be due to similar substratum and water velocity preferences, although at both the regional and microhabitat scales E. berilloni prefers wider streams than G. duebeni celticus . This study reveals a decline in the endangered G. duebeni celticus since 1970.
Main conclusions  The longitudinal and local distributions of G. duebeni celticus , and the higher-than-expected co-occurrence of the species with G. pulex , suggest that the decline of the endemic species may be due to changes in the environment and/or interference from native G. pulex , which is expanding its range in Brittany. The results are discussed as regards to the consequences for regional biodiversity.