Different ammonia tolerances may facilitate spatial coexistence of Gammarus roeselii and the strong invader Dikerogammarus villosus

The introduction of invasive species that can replace native species is one of the most critical threats to the biodiversity of aquatic systems. Here we investigated the potential contribution of one factor to the coexistence of the indigenous amphipod Gammarus roeselii and the invasive amphipod Dikerogammarus villosus in the same ecosystem (Lake Constance) within different microhabitats. We quantitatively studied the influence of ambient ammonia concentrations on the distributions of the two amphipod species. We also assessed the ammonia tolerance ranges of both species in laboratory experiments by measuring mortality rate, precopula disruption, egg mortality, and microhabitat choice. The proportion of G. roeselii among the two amphipod species was significantly positively related to the ammonia concentration in the water, which indicated that the distribution of the invasive D. villosus was limited at high ammonia concentrations. Although the mortality rates of the two species did not significantly differ, G. roeselii was more tolerant to ammonia with regard to precopula disruption, egg mortality, and microhabitat choice. The effective ammonia concentrations that led to a significantly reduced direct reproductive success in D. villosus were within the range of the highest field concentrations measured, where only G. roeselii occurred. D. villosus may have a smaller range than the indigenous G. roeselii partially because of its lower tolerance to higher ammonia concentrations, which lead to reduced reproductive success. Beside other habitat parameters differences on ammonia tolerance between the two amphipods might allow their coexistence along a gradient of microhabitats in Lake Constance.     

Growth and reproductive characteristics of rarely observed resident female Dolly Varden (Salvelinus malma malma) in North America

The replacement of native species by invasive species is one of the most critical threats to the biodiversity of aquatic systems today. However, little is known about potential effects of species invasions on the genetic diversity of indigenous species in cases where the latter coexist with the invader. Here we present an example of the indigenous amphipod Gammarus roeselii, which has been partly replaced by the invasive Dikerogammarus villosus in Lake Constance (C-Europe) and now mostly exists in small, isolated populations. We compared the genetic diversity, population structure, indicators for bottlenecks and migrations rates of G. roeselii before and after the invasion event from samples collected between 1999 and 2013. We expected a genetic impoverishment in the reduced and segregated populations of G. roeselii. However, no genetic measure on G. roeselii differed temporally or spatially over the investigated period, which indicates that D. villosus has not yet had an impact on G. roeselii at the genetic scale. Hence, even though a decline in population size of G. roeselii was found in Lake Constance, our results on the genetic scale contribute to recent findings that the overall impact of D. villosus on native species is not as strong as often discussed.

     

Zebra mussels mediate benthic–pelagic coupling by biodeposition and changing detrital stoichiometry

1. The zebra mussel ( Dreissena polymorpha ) is one of the most successful invasive species; it has colonised many aquatic systems in Europe and North America with strong impacts on various ecosystem processes. The effect of D. polymorpha filtration on pelagic seston concentrations has been quantified in several studies, but the magnitude and stoichiometry of the transfer of sestonic biomass into benthic detritus by D. polymorpha and the accompanying enrichment of the benthic habitat is still under-investigated.
2. We studied biodeposition by zebra mussels in two series of laboratory experiments with the food algae Cryptomonas erosa and Scenedesmus obliquus . We also measured the year-round biodeposition rate under natural conditions in the oligotrophic Lake Constance.
3. In all experiments, zebra mussel biodeposition was linearly related to seston concentration. In the field, the relationship changed with a seasonal shift in algal composition and lower biodeposition rates during the spring algal bloom.
4. For both algal species in laboratory experiments, biodeposited material was depleted in phosphorous at an algal concentration ≤0.6 mg ash-free dry mass L⁻¹, but not at higher concentrations. This effect was not observed in the field, probably because of high variation in C : N : P stoichiometry.
5. By mediating the transfer of pelagic resources into the benthos zebra mussels provide a sufficient amount of detritus for benthic invertebrates, especially during summer. Thus, material biodeposited by the mussels might increase benthic secondary production from pelagic resources, and zebra mussels are important mediators of this flux of organic matter from the pelagic zone into the benthos.     

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.     

Consequences of the colonisation of leaves by fungi and oomycetes for leaf consumption by a gammarid shredder

1. Leaf litter breakdown by shredders in the field is affected by leaf toughness, nutritional value and the presence of secondary compounds such as polyphenols. However, experiments involving the use of single fungal strains have not supported the assumption that leaf parameters determine food selection by shredders perhaps because of a failure to test for high consumption prior to isolation of fungal strains, overrepresentation of hyphomycetes or the potential effects of accompanying bacteria. In this study, we used bacteria‐free, actively growing fungi and oomycetes isolated from conditioned leaf litter for which a shredder had already shown high consumption rates. 2. Black alder (Alnus glutinosa) leaf litter was exposed to the littoral zone of Lake Constance in autumn, and subsamples were analysed for leaf parameters and consumption by Gammarus roeselii under standard conditions at regular intervals. On dates with a high consumption rate of the exposed leaves, 14 single strains of fungi and oomycetes were isolated, freed of bacteria and grown on autoclaved leaves. 3. Six of eight measured leaf parameters of exposed leaves were significantly correlated with Gammarus consumption rates, with high colinearity among leaf parameters hampering the identification of causal relations between leaf parameters and feeding activity. 4. When single strains of fungi and oomycetes were grown on autoclaved leaf litter, toughness of colonised leaves was always lower than in the control and the content of protein, N and P were increased. There were pronounced strain‐specific effects on leaf parameters. Consumption rates also differed significantly, with nine of fourteen isolates consumed at higher rates than controls and none proving to be a deterrent. Protein and polyphenol content were significantly correlated with consumption rates. Oomycete‐colonised leaves were consumed at similar rates but were of lower food quality than fungi‐colonised leaves. 5. We argue that direct strain‐specific attractant or repellent effects of fungi and oomycetes on consumption by G. roeselii are not important. However, we found indirect strain‐specific role operating via effects on leaf parameters.     

The very hungry amphipod: the invasive <Emphasis Type="Italic">Dikerogammarus villosus</Emphasis> shows high consumption rates for two food sources and independent of predator cues

The invasion of the Ponto–Caspian amphipod Dikerogammarus villosus in European rivers is assumed to reduce macroinvertebrate diversity and to alter ecosystem functions. D. villosus shows an extraordinarily flexible feeding behavior including the ability to use various food sources. On the other hand, its response to predation risk seems to depend on environmental factors. To evaluate the ecological function of D. villosus, we estimated the daily food consumption for different food sources and analyzed potential effects of predator avoidance behavior on feeding. D. villosus consumption of willow leaves or chironomid larvae was quantified in 24-h laboratory experiments with and without kairomones of the European bullhead (Cottus gobio). Consumption rates were estimated based on gut content and gut evacuation rate under semi-natural laboratory conditions enabling the animals to feed over the whole time of the evacuation rate experiment. We observed very high evacuation rates and consequently high consumption rates up to 89% of body weight per day. Consumption rates differed significantly between food sources: D. villosus ingested more leaves than chironomid larvae. In contrast, predator cues did not affect the feeding of D. villosus. This might be explained by its strong refuge affinity and probably benefits its successful invasion. A comparison of the estimated consumption rates with results of an own consumption experiment (and other studies) under more artificial conditions indicated that more natural conditions result in higher consumption rates. Consequently, feeding rates from highly artificial experiments should be used with great caution to assess the ecosystem function of D. villosus.     

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.     

Colonization patterns and impacts of the invasive amphipods Chelicorophium curvispinum and Dikerogammarus villosus in the IJsselmeer area, The Netherlands

During the 1980s, Chelicorophium curvispinum and Dikerogammarus villosus colonized the lakes of the IJsselmeer area in The Netherlands. With the arrival of C. curvispinum the indigenous Apocorophium lacustre retreated to a small, sheltered bay with particular microconditions. The arrival of D. villosus is probably linked to a decrease of Gammarus tigrinus and possibly the disappearance of the gastropod Theodoxus fluviatilis. However, the occurrence of D. villosus may be restricted to stones along the banks of the lakes, and even there, G. tigrinus seems to remain stable at lower population densities. In samples from the beds of Dreissena polymorpha on the bottom of the lakes, D. villosus was virtually absent and densities of C. curvispinum were relatively low. G. tigrinus was here the single gammaridean species. Other changes in densities of invertebrates, like strong decreases in densities of Tricladida, Hirudinea, Asellidae, D. polymorpha and several gastropods, differ in details of timing from the arrival of the invaders, and are at least partly linked to other factors. In the River IJssel, the main source of invaders to the lakes, similar decreases in species abundance followed exceptionally high discharge events, but preceded both a relatively strong decrease of chlorophyll a levels and the arrival of D. villosus. This suggests that this arrival was not a direct cause of the recorded declines, although it may have prevented recovery.     

Effect of leaf litter characteristics on leaf conditioning and on consumption by Gammarus pulex

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The shredding activity of gammarids facilitates the processing of organic matter by the subterranean amphipod Niphargus rhenorhodanensis

1. The functional feeding group approach has been widely used to describe the community structure of benthic invertebrates in relation to organic matter resources. Based on this functional framework, positive interactions between feeding groups (especially shredders and collector‐gatherers) were postulated in the River Continuum Concept. However, relationships with organic matter have been poorly documented for invertebrates living in the hyporheic zone. 2. We hypothesised that the common subterranean amphipod Niphargus rhenorhodanensis would feed on fine particulate organic matter (FPOM), which is more abundant than coarse particulate organic matter (CPOM) in hyporheic habitats, and should be favoured by the occurrence of shredders that produce FPOM from CPOM. 3. We used laboratory experiments to quantify leaf litter processing by N. rhenorhodanensis and a common shredder, the surface amphipod Gammarus roeselii. We estimated rates of feeding and assimilation (using nitrogen stable isotopes) of the two species separately and together to reveal any potential shredder–collector facilitation between them. 4. Measured leaf litter mass loss showed that N. rhenorhodanensis did not act as a shredder, unlike G. roeselii. Organic matter dynamics and ¹⁵N/¹⁴N ratios in tissues of niphargids indicated that N. rhenorhodanensis was a collector‐gatherer feeding preferentially on FPOM. We also found a positive influence of the gammarid shredders on the assimilation rate of N. rhenorhodanensis, which fed on FPOM produced by the shredders, supporting the hypothesis of a positive interaction between surface shredders and hyporheic collector‐gatherers.     

A local gammarid uses kairomones to avoid predation by the invasive crustaceans Dikerogammarus villosus and Orconectes limosus

Since the 1980s the Ponto-Caspian gammarid Dikerogammarus villosus has spread throughout Europe while displacing native species and is predicted to invade further continents. After it was introduced into Europe in the 1890s the North American crayfish Orconectes limosus spread throughout Europe and served as a vector to displace native crayfish as well. In Lake Constance (Germany) the previously dominant gammarid Gammarus roeselii is subjected to both of these invasive crustaceans. In our experiments both species placed predation pressure on G. roeselii. Kairomone perception tests in a Y-maze revealed the capability of the gammarids G. roeselii and D. villosus to perceive and avoid the scent of the predator crayfish O. limosus. Both species also avoided the kairomone of the other gammarid, but did not avoid the scent of their own species. This taxa specific behavior suggests that taxa specific signals are used. This behavior can help the gammarids avoid shelters previously occupied by predators.     

Assimilation efficiency of adult Kittiwakes and Brünnich's Guillemots fed Capelin and Arctic Cod

The mean assimilation efficiencies of 10 adult Kittiwakes (Rissa tridactyla) and 10 Brünnich's Guillemots (Uria lomvia) fed on Capelin (Mallotus villosus) were 77.5% and 74.4%, respectively. When fed on Arctic Cod (Boreogadus saida) they were 83.1% and 78.2%, respectively. After correction for nitrogen retention, the assimilation efficiencies decreased to 72.2%, 70.6%, 81.2% and 74.7%, respectively. Kittiwakes and Brünnich's Guillemots seem to have the same ability to utilize the energy of the different food items. The differences in assimilation efficiencies when fed two fish species was mainly related to the fat content of the fish.     

Ontogenetic shift in the trophic role of the invasive killer shrimp Dikerogammarus villosus: a stable isotope study

The introduction of the amphipod Dikerogammarus villosus in European fresh waters is to date recognized as a threat to the integrity of invaded communities. Predation by D. villosus on native benthic invertebrates is assumed as the key determinant of its ecological impact, yet available information describe the species as a primary consumer as well as a carnivore depending on local conditions. Here, we assessed the trophic position (TP) of D. villosus in Lake Trasimeno, a recently invaded lentic system in central Italy, using the CN isotopic signatures of individuals captured in winter spanning two orders of magnitude in body size. TP estimations were compared with those characterizing the native amphipod Echinogammarus veneris and other representative invertebrate predators. On average, D. villosus showed a trophic position higher than E. veneris, and comparable with that of odonate nymphs. An in-depth analysis revealed that large-sized individuals had a trophic position of 3.07, higher than odonates and close to that of the hirudinean predator Erpobdella octoculata, while small-sized specimens had a trophic position of 2.57, similar to that of E. veneris (2.41). These findings indicate that size-related ontogenetic shifts in dietary habits may per se vary the nature of the interaction between Dikerogammarus villosus and native invertebrates from competition to predation. Information collated from published isotopic studies corroborated the generality of our results. We conclude that intra-specific trophic flexibility may potentially amplify and make more multifaceted the impact of the species on other invertebrate species in invaded food webs.

     

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.     

Size matters: predation of fish eggs and larvae by native and invasive amphipods

Invasive predators can have dramatic impacts on invaded communities. Extreme declines in macroinvertebrate populations often follow killer shrimp (Dikerogammarus villosus) invasions. There are concerns over similar impacts on fish through predation of eggs and larvae, but these remain poorly quantified. We compare the predatory impact of invasive and native amphipods (D. villosus and Gammarus pulex) on fish eggs and larvae (ghost carp Cyprinus carpio and brown trout Salmo trutta) in the laboratory. We use size-matched amphipods, as well as larger D. villosus reflecting natural sizes. We quantify functional responses, and electivity amongst eggs or larvae and alternative food items (invertebrate, plant and decaying leaf). D. villosus, especially large individuals, were more likely than G. pulex to kill trout larvae. However, the magnitude of predation was low (seldom more than one larva killed over 48 h). Trout eggs were very rarely killed. In contrast, carp eggs and larvae were readily killed and consumed by all amphipod groups. Large D. villosus had maximum feeding rates 1.6–2.0 times higher than the smaller amphipods, whose functional responses did not differ. In electivity experiments with carp eggs, large D. villosus consumed the most eggs and the most food in total. However, in experiments with larvae, consumption did not differ between amphipod groups. Overall, our data suggest D. villosus will have a greater predatory impact on fish populations than G. pulex, primarily due to its larger size. Higher invader abundance could amplify this difference. The additional predatory pressure could reduce recruitment into fish populations.     

Differential effect of wave stress on the physiology and behaviour of native versus non-native benthic invertebrates

In fresh waters, non-native invertebrate species preferentially spread via navigation waterways, where they often dominate assemblages. Littoral habitats in navigation waterways are regularly exposed to ship-induced waves. We conducted experiments to test the effects of artificial wave exposure on the relative performance of wide-spread native and non-native species. We compared physiological and behavioural response variables (i.e. growth rate, glycogen content, feeding and swimming activity) of two amphipods (native Gammarus roeselii and non-native Dikerogammarus villosus) and two gastropods (native Bithynia tentaculata and non-native Physella acuta) subject to wave and control (i.e. no wave) treatment flumes across a 6-week period. Growth, and in part glycogen content (as a measure of energy storage), were significantly reduced after exposure to waves in native invertebrates, but not in non-native invertebrates. The reduction in growth may be associated with the disturbance effects of waves, such as the higher swimming activity of G. roeselii and lower food uptake of B. tentaculata. In comparison, the effective hiding behaviour observed for D. villosus and good swimming ability of P. acuta, were identified as important traits facilitating the successful colonisation of the harsh habitat conditions of littoral waterways. Our study demonstrates that artificial wave regimes may contribute significant selective pressure, thus explaining the observed dominance of non-native species in navigational waterways. The success of non-native species under the harsh hydraulic habitat conditions of these socio-economically driven ecosystems may consequently be traced directly to behavioural and/or physiological traits.     

Attraction of the amphipod <Emphasis Type="Italic">Gammarus pulex</Emphasis> to water-borne cues of food

We examined the ability of the amphipod Gammarus pulexto detect chemical cues released from potential food sources. Therefore, response of G. pulex to chemical cues from food was tested in paired-choice laboratory experiments. Comparisons were made between artificial and natural leaves, with and without the importance of aufwuchs, and with different components of the aufwuchs community. Our study demonstrated that G. pulex actively chose its food and that G. pulex is most strongly attracted to the aufwuchs on discs rather than to the leaf itself. Fungi and bacteria are more important in the food selection process than algae probably because fungal and bacterial cues are more specific cues for decaying leaves than algal cues, since algae also grow on mineral substrates and then do not contribute to leaf decomposition.     

Importance of fungi in the diet of Gammarus pulex and Asellus aquaticus

An important component of the interaction between macroinvertebrates and leaf litter in streams in the extent to which consumers can differentiate between undecomposed and decomposing leaves. The detritivores Gammarus pulex and Asellus aquaticus fed preferentially on conditioned rather on unconditioned leaf material. Growth in A. aquaticus was significantly reduced when unconditioned leaves were provided, but in G. pulex no significant effect of conditioning on growth was observed. The capacity of G. pulex to tolerate reductions in food quality seems to be a consequence of a compensatory system in which respiration rates change to compensate for reductions in food quality. In this way a constant growth rate is maintained. Increases in ingestion rates to compensate for low quality food were not observed.     

Molecular markers for systematic identification and population genetics of the invasive Ponto‐Caspian freshwater gammarid Dikerogammarus villosus (Crustacea,Amphipoda)

The Ponto‐Caspian amphipod, Dikerogammarus villosus, is an invasive species of many European rivers. First, we show that size difference of nrDNA ITS1 allows discriminating D. villosus from Dikerogammarus bispinosus, a closely related but morphologically hardly distinguishable species. Second, we present two types of polymorphic markers for D. villosus, three microsatellites and two single nucleotide polymorphisms (SNPs) of mtDNA COI gene, which were scored by polymerase chain reaction‐single strand conformational polymorphism (PCR‐SSCP). These markers will be very useful in studying population genetics of D. villosus.     

Nutrients are assimilated efficiently by Lymantria dispar caterpillars from the mature leaves of trees in the Salicaceae

The efficient aquisition of nutrients from leaves by insect herbivores increases their nutrient assimilation rates and overall fitness. Caterpillars of the gypsy moth (Lymantria dispar L.) have high protein assimilation efficiencies (PAE) from the immature leaves of trees such as red oak (Quercus rubra) and sugar maple (Acer saccharum) (71–81%) but significantly lower PAE from their mature leaves (45–52%). By contrast to this pattern, both PAE and carbohydrate assimilation efficiencies (CAE) remain high for L. dispar larvae on the mature leaves of poplar (Populus alba × Populus tremula) grown in greenhouse conditions. The present study tests two alternative hypotheses: (i) outdoor environmental stresses cause decreased nutrient assimilation efficiencies from mature poplar leaves and (ii) nutrients in the mature leaves of trees in the poplar family (Salicaceae) remain readily available for L. dispar larvae. When poplar trees are grown in ambient outdoor conditions, PAE and CAE remain high (approximately 75% and 78%, respectively) in L. dispar larvae, in contrast to the first hypothesis. When larvae feed on the mature leaves of species in the Salicaceae [aspen (Populus tremuloides), cottonwood (Populus deltoides), willow (Salix nigra) and poplar], PAE and CAE also remain high (68–76% and 72–92%, respectively), consistent with the second hypothesis. Larval growth rates are strongly associated with protein assimilation rates, and more strongly associated with protein assimilation rates than with carbohydrate assimilation rates. It is concluded that tree species in the Salicaceae are relatively high‐quality host plants for L. dispar larvae, in part, because nutrients in their mature leaves remain readily available.