Mitochondrial DNA polymorphism among alpine populations of Gammarus lacustris (Crustacea, Amphipoda)

1. Mitochondrial DNA sequence analysis was used to investigate the phylogeographic structure among populations of the amphipod Gammarus lacustris isolated in high altitude lakes from the Northern to the Southern Alps, and to investigate the historical and demographic events that may have accompanied this differentiation.
2. The comparison of a 376‐base segment of the mitochondrial cytochrome c oxidase I gene in fifty‐four specimens from nine different populations reveals no variation within populations. However, northern populations appear divergent from those from the Southern Alps. A particular population, located at the limit between the Northern and Southern Alps shows an intermediate haplotype related to the southern lineage.
3. Consideration of the different mitochondrial lineages, together with geographical differences among populations, suggests a scenario for the post‐glacial colonization of G. lacustris in the alpine lakes.     

Parasite altered micro-distribution of Gammarus pulex (Crustacea: Amphipoda)

In a river survey, Gammarus pulex amphipods both unparasitised and parasitised with the acanthocephalan Echinorhynchus truttae were distributed similarly with respect to flow regimen, tending to be more abundant in faster, shallower, riffle patches. However, there was a higher prevalence of parasitism in faster, shallower areas than in slower, deeper areas and abundance correlated with macrophyte coverage for unparasitised but not parasitised amphipods, indicating subtle differences in habitat usage. A laboratory 'patch' simulation indicated that parasitism influenced micro-distribution. There were higher proportions of unparasitised amphipods in/under stone substrates and within weed. In contrast, there were higher proportions of parasitised amphipods in the water column and at the water surface. As the experiment progressed, unparasitised but not parasitised amphipod habitat usage shifted from those micro-habitats above the substrate and in the water column to those in/under the substrates. Experiments also demonstrated that parasitised amphipods were more active and had a greater preference for illumination. Previous studies of the effects of acanthocephalan parasitism of amphipod hosts have focussed on how drift behaviour is altered, now we show that subtle differences in micro-habitat usage could translate to greatly increased vulnerability to fish predation. We discuss how aggregation of parasitised individuals within specific habitats could promote parasite transmission.     

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.     

Distribution of Gammarus lacustris Sars (Amphipoda, Gammaridae) in Lake Shira (Khakasia, Siberia) and laboratory study of its growth characteristics

Spatial distribution of Gammarus lacustris in Lake Shira and growth of young specimens that feed on lake biota were studied. The amphipods have been shown to inhabit the littoral, sublittoral and the upper aphytal zones of the lake on stony-sandy soil and silted sand. The young and adult individuals stay apart. The young live in submerged or semi-submerged vegetation in the littoral, the adults in the sublittoral and upper aphytal zones. Maximum density of amphipods was encountered in the areas influenced by human activity.The feeding experiments revealed that the lake plankton is a more important food source for G. lacustris than any other food species. The specific growth rate measured was 0.039 d^–1, with a length increment 0.095 mm d^–1.     

Field estimates of feeding rate for Gammarus pseudolimnaeus (Crustacea: Amphipoda) in the Credit River, Ontario

SUMMARY. The feeding rate of G. pseudolimnaeus was measured monthly for 7 months in the field by monitoring the decline in weight of gut contents when the amphipod was starved. This decline was modelled by an exponential regression of weight on time. As the amphipods appeared to be continuous feeders, feeding rate was calculated by multiplying the dry weight of a full gut by the specific rate of emptying, i.e., the slope of the exponential regression. Specific rate of emptying was independent of animal size, but increased with temperature. Therefore, food has a longer period in which to be digested at low temperatures, which suggests that assimilation efficiency may increase.
However, the assimilation efficiency of amphipods feeding on decaying maple leaves in the laboratory was only 10% and did not vary with temperature. Ingestion and egestion rates were measured in the laboratory by weighing amounts eaten and defecated. The turnover time of the contents of a full gut in the laboratory often agreed very well with turnover time measured in the field, i.e., the reciprocal of the specific rate of emptying, thus confirming the use of an exponential regression.     

The ultrastructure of the sinus gland of Gammarus oceanicus (Crustacea: Amphipoda)

Summary The sinus gland of Gammarus oceanicus, like that of other crustaceans, is composed of three elements: neurosecretory axons, glial cells and stromal sheath. Five neurosecretory axon types are identified on the basis of granule diameter, shape, and electron density, and axon matrix density. Exocytosis appears to be the major release mechanism of neurosecretory material. The preterminal regions of neurosecretory axons contain axoplasmic reticulum and neurotubules. Their arrangement in the axon and relationship with one another suggest a transport function. Multilamellar bodies are found in the terminal regions of neurosecretory axons. They arise from mitochondria and may be involved in granulolysis.The technical assistance of G.A. Bance, statistical assistance of D. MacCharles and D.W. Hagen, and financial support provided by the University of New Brunswick Research Fund to K.H. are gratefully acknowledged     

Anatomy of the extrinsic gut musculature of Gammarus minus (Crustacea: Amphipoda)

Inserting on the buccal and esophageal foregut of Gammarus minus are numerous pairs of serially arranged dorsal dilator muscles, a single pair of lateral muscles, and two pairs of posterior muscles. Muscles of the cardiac stomach include three dorsal sets, a single pair associated with the pterocardiac ossicles, and two pairs inserting on the ventral aspect. A single pair of muscles inserts on the lateral aspect of the pyloric stomach. The extrinsic muscles of the foregut originate from exoskeletal apodemes of the cephalothoracic cuticle, sockets of the mandible, and a maxillary bridge that lies just ventral to the cardiac stomach. The extrinsic musculature of the hindgut is restricted to the rectal region and consists of paired dorsal and ventral series in an X-configuration. A single unpaired muscle inserts on the ventral midline. Extrinsic muscles of the hindgut originate from the integument of the last pleonic segment. The general arrangement of extrinsic gut muscles in G. minus is similar but not identical to that of other amphipods studied. However, the pattern is quite different from that of other malacostracans.     

The influence of experimental scale on estimating the predation rate of Gammarus lacustris (Crustacea: Amphipoda) on Daphnia in an alpine lake

We examined the effect of experimental scale on the predationrate of Gammarus lacustris preying on Daphnia middendorffianain 20 l cubitainers, 2800 l mesocosms and diurnal whole-lakesurveys. The predation rate differed significantly among experimentalscales. It was highest in the 20 l cubitainers, intermediatein the 2800 l mesocosms and lowest in the lake. At all experimentalscales, the predation of G.lacustris on D.middendorffiana increasedas a linear function of prey density (Type I functional response).Very different predation rates were calculated for the wholelake depending on whether D.middendorffiana densities were estimatedfrom whole-water column hauls or stratified depth hauls. Thishighlights the importance of accurately determining the spatialdistribution of prey for estimating reliable in situ predationrates. We also determined prey selection by G.lacustris. Daphniamiddendorffiana was strongly selected over the calanoid copepodHesperodiaptomus arcticus. When presented with different sizedDaphnia, G.lacustris preyed size selectively on individuals2 mm. Our results show that the predation rate of small invertebratepredators is sensitive to a wide range of experimental conditionsand needs to be corroborated at a variety of scales. We concludethat the impact of G.lacustris on zooplankton communities wouldbe strongest in small fishless lakes or ponds.     

Identifying Chinese species of Gammarus （Crustacea： Amphipoda） using DNA barcoding

Using a standard cytochrome c oxidase I sequence, DNA barcoding has been shown to be effective to distinguish known species and to discover cryptic species. Here we assessed the efficiency of DNA barcoding for the amphipod genus Gammarus from China. The maximum intraspecific divergence for widespread species, Gammarus lacustris, was 3.5%, and mean interspecific divergence reached 21.9%. We presented a conservative benchmark for determining provisional species using maximum intraspecific divergence of Gammarus lacustris. Thirty-one species possessed distinct barcode clusters. Two species were comprised of highly divergent clades with strong neighbor-joining bootstrap values, and likely indicated the presence of cryptic species. Although DNA barcoding is effective, future identification of species of Gammarus should incorporate DNA barcoding and morphological detection[Current Zoology 55(2):158-164,2009].     

The dynamics of predation on Gammarus spp. (Crustacea: Amphipoda)

Gammarus spp. (Crustacea: Amphipoda) are widespread throughout a diverse range of marine, freshwater and estuarine/brackish habitats, often dominating benthic macroinvertebrate communities in terms of both numbers and/or biomass. Gammarus spp. are the dominant macroinvertebrate prey items of many fish, whether as a seasonal food source or a year-round staple. Selective predation by fish on Gammarus spp. is often linked to parasitism and the body size of the prey. Gammarus spp. populations are under increasing threat from both pollution and replacement/displacement by introduced species. Loss of populations and species invasions/replacements could have significant impacts on native predator species if the predator(s) cannot successfully adapt their feeding patterns to cope with non-indigenous Gammarus prey species. Despite this, many fish predation studies do not identify Gammarus prey to species level. This lack of precision could be important, as Gammarus spp. exhibit wide variations in physiochemical tolerances, habitat requirements, abilities to invade and susceptibility to replacement. Although rarely acknowledged, the impacts of nonpiscean predators (particularly macroinvertebrates) on Gammarus prey species may frequently be stronger than those exerted by fish. A major aim of this review is to ascertain the current importance of Gammarus as a prey species, such that the implications of changes in Gammarus spp. populations can be more accurately assessed by interested groups such as ecologists and fisheries managers. We also review the dynamics of Gammarus spp. as prey to a diverse array of mammals, birds, amphibians, insects, flatworms, other crustaceans such as crabs and crayfish and, perhaps most importantly, other Gammarus spp.     

Relation between habitat and antennular segmentation in Gammarus and Echinogammarus (Crustacea: Amphipoda)

In 140 species of the amphipod genera Gammarus and Echinogammarus the ratio was calculated between the number of segments of the accessory and main antennular flagellum. Minor inter-and intrapopulational variation of this ratio was found in four species of the genus Gammarus. Seasonal variation was studied in one species of this genus. For both genera a clear correlation was found between antennular composition and the occurrence in marine or freshwater habitats.     

Ecology of <Emphasis Type="Italic">Gammarus lacustris</Emphasis> Sars (Crustacea: Amphipoda) in Transbaikalian water bodies

The data on Gammarus lacustris Sars, 1863 ecology in Transbaikalian water bodies was integrated by the example of Lakes Khalanda, Bain-Tsagan, Arakhlei, Zun-Torei, and Bain-Bulak. The impact of the major environmental factor on the pattern of G. lacustris spatial distribution, linear size, growth rate, time of reaching sexual maturity, and number of reproduced generations was shown. The role of G. lacustris in the ecosystem of studied water bodies was evaluated and the adaptive significance of the revealed life strategies was considered.     

Molecular evidence for further overlooked species within the Gammarus fossarum complex (Crustacea: Amphipoda)

The freshwater amphipod Gammarus fossarum Koch, in Panzer, 1836 is a locally abundant keystone species mainly occuring in European headwaters but also in larger rivers. Genetic studies in the past 25 years have revealed three cryptic species within nominal G. fossarum (types A, B and C). Assignments of specimens to these types were based on allozyme and 16S markers. Today, a fragment of the cytochrome c oxidase subunit 1 (CO1) is primarily used as a genetic marker for species assignments (‘DNA Barcoding’), yet not a single CO1 sequence of G. fossarum is available in the Barcode of Life Database. We analysed new CO1 and 16S data for German, Hungarian and Croatian G. fossarum specimens and compared them with 16S and CO1 sequences of G. fossarum from GenBank. Thereby, we close the gap between traditional allozyme- and 16S-based species assignments and modern CO1 barcoding. Studying genetic variation in 55 specimens from 29 populations, we identified between 11 and 23 novel and genetically distinct clades using distance- and tree-based methods. Our results suggest that G. fossarum comprises several additional, yet unrecognised, species in particular from the Balkan region. Therefore, a taxonomic revision and biogeographic reconsideration of the G. fossarum complex is urgently needed.     

Differential drift and parasitism in invading and native Gammarus spp. (Crustacea: Amphipoda)

Invading and native species often interact directly, such as by predation, producing patterns of exclusion and coexistence. Less direct factors, such as interactions with the broader abiotic and biotic environment, may also contribute to such patterns, but these have received less recognition. In Northern Ireland, the North American Gammarus tigrinus has invaded freshwaters populated with the native Gammarus duebeni celticus , with intraguild predation between the two implicated in their relative success. However, these species also engage in day and night "drifting", an activity that subjects amphipods to intense predation from fish and wildfowl. Sampling of two rivers where the invader and native co-occur showed that, compared with the benthos, G. tigrinus was underrepresented and G. d. celticus overrepresented in the drift. In addition, G. tigrinus were free from parasites, whereas some G. d. celticus harboured the acanthocephalans Polymorphus minutus and Echinorhynchus truttae and the muscle wasting microsporidian Pleistophora sp. (new species). Compared with the benthos, G. d. celticus parasitized with P. minutus were overrepresented and unparasitized individuals underrepresented in the drift. The opposite pattern was found with Pleistophora sp . In laboratory experiments, G. tigrinus were less positively phototropic and less "active" than G. d. celticus (unparasitized animals). Polymorphus minutus increased G. d. celticus positive phototropism and activity, while Pleistophora sp. increased positive phototropism but decreased activity. Previous studies show that the invader G. tigrinus is more disadvantaged by intraguild predation from the native G. d. celticus than vice versa. However, the native appears more disadvantaged with respect to drift, parasitism and the interaction of the two. These factors may mitigate direct interactions and help explain complex patterns of coexistence between these invader and native species.     

Quantitative aspects of oxygen uptake by Gammarus (Crustacea, Amphipoda): a critical review

SUMMARY. Gill structure and function are brietly considered, as are the main factors affecting the oxygen content of water. The review deals with quantitative, analytical aspects of oxygen uptake related to body size, temperature, oxygen content of water, salinity, water velocity, substratum and activity of Cammarus. A theoretical model relates oxygen uptake to growth and life-span of individuals. Several aspects of regression analysis, as applied to measurements of oxygen uptake, are examined and constructively criticized. The energetic cost of osmoregulation is calculated for G. pulex. The review concentrates on freshwater species of Gammarus but work on brackish-marine species is also included. A more detailed summary and conclusions are given at the end of the review.     

Isolation and characterization of the first eight microsatellite loci in Gammarus fossarum (Crustacea, Amphipoda) and cross-amplification in Gammarus pulex and Gammarus orinos

We characterized the first microsatellite markers for Gammarus fossarum. Eight loci gave satisfactory amplification patterns in two stream populations (Southern France) with number of alleles ranging from 2 to 10 and expected heterozygosity from 0.076 to 0.857. We performed cross-amplification in two closely related gammarid species, Gammarus pulex and Gammarus orinos. Among the eight tested microsatellite loci, four correctly amplified in G. pulex and three in G. orinos.