Effects of coinfection with Pomphorhynchus bulbocolli on development of leptorhynchoides thecatus (Acanthocephala) in amphipods (Hyalella azteca)

The effect of the presence of Pomphorhynchus bulbocolli on the development of Leptorhynchoides thecatus (Acanthocephala) in amphipod intermediate hosts (Hyalella azteca) was examined. Two groups of amphipods were exposed either to eggs of both species (experimental) or only to L. thecatus eggs (control). Amphipods of both groups were dissected 39 days postexposure. The percentage of L. thecatus at the cystacanth stage and mean abundance of cystacanths and precystacanths per amphipod were determined. Contingency table analysis and Fisher's exact tests demonstrated that a significantly smaller proportion of L. thecatus reached the cystacanth stage in coinfected amphipods of the experimental group than either worms in the control group or worms in L. thecatus-only infections of the experimental group. Interspecific interaction between acanthocephalans in intermediate hosts has not been reported previously. Coinfection reduces the number of L. thecatus in amphipods available for transmission to definitive hosts. Habitat separation by eggs of L. thecatus and P. bulbocolli might reduce this effect if amphipods are less likely to become coinfected when eggs are in different microhabitats than when they are not.     

Drift and upstream movement of invertebrates in a springbrook community ecosystem

Structure, drift, and upstream movement of populations of benthic macroinvertebrates, in particular Synurella dentata Hubricht and Lirceus fontinalis Raf., were examined within a temperate spring ecosystem. Chemical and physical aspects of the springbrook were also investigated and life histories of the gammarids and asellids noted.Chemically and physically the spring proved both constant and predictable, much more so than other lotic systems.Species diversity was low from November through February and increased in March, April, and May. Equitability followed the same trends as species diversity. Both indices were most affected by large fluctuations in the populations of aquatic insects.Significant changes in the numbers of amphipods, isopods, and total macroinvertebrates was evident over a seven month period. Males were present in the isopod population year-round, but only from November to January in the amphipod population. Breeding by the isopods occurred throughout the year and peaked during winter. Amphipods copulated only in the late fall and early winter.Significant diel peaks in the amphipod, isopod, and total invertebrate drift negatively correlated with light intensity levels. Amphipods and isopods did not exhibit any preferential upstream movement during either the day or night; however, total macroinvertebrate upstream movement was greater at night. The total number of invertebrates moving upstream were lower than values reported from other lotic environments.     

Movement of Pomphorhynchus bulbocolli larvae from the hemocoel to the peripheral circulation of Gammarus pseudolimnaeus

Acanthors and early acanthellae of Pomphorhynchus bulbocolli were observed in the peripheral circulation of experimentally infected laboratory-reared Gammarus pseudolimnaeus. This has not previously been reported for larval P. bulbocolli during development in the intermediate host. The timing of larval movement from hemocoel to vessels and sinuses was related to intensity of infection. In an experimental group with a low mean intensity of infection, the movement of larval P. bulbocolli into the peripheral circulation occurred later (9 days postfeeding) than in a group with a high mean intensity of infection (2 days postfeeding). Larvae were observed in the peripheral circulation for a longer period of time in amphipods from the group with the high mean intensity of infection. Dead and slightly melanized acanthors and early acanthellae were present in the hemocoel at the time living larvae moved into the peripheral circulation. Only 1 incidence of a hemocytic response to the larvae was observed in the first 20 days postfeeding.     

Seasonal distribution of sympagic amphipods near Chesterfield Inlet, N.W.T., Canada

The seasonal distribution of sympagic amphipods was investigated in the Chesterfield Inlet area of northwestern Hudson Bay (63°30′N). Amphipod abundance was measured by photographic samples and species composition was determined by sweep net samples. Twelve species of amphipods were collected, the most common being Ischyrocerus anguipes, Pontogeneia inermis, Apherusa megalops and Weyprechtia pinguis. The major environmental variable affecting amphipod distribution was water depth. Amphipod abundance was highest near 20 m and near zero past 50 m. The maximum recorded abundance was 1367 m⁻². A minor factor affecting the distribution of amphipods was snow depth, through its modifying effect on light and thereby the growth of ice algae. Amphipods began to inhabit the sea ice shortly after its formation. From the beginning of March, the number of amphipods on the ice increased steadily to about the 3rd week of April, after which numbers declined. This pattern coincided with the seasonal ice algae abundance. Amphipods reduced ice algal biomass over 20-m depth by 63%. No evidence of diurnal changes in abundance was observed. Received: 15 May 1996 / Accepted: 4 November 1996     

Effect of cystacanth body size on adult success

Laboratory-reared cystacanths of Leptorhynchoides thecatus (Acanthocephala: Rhadinorhynchidae) were used to study the effect of cystacanth size on adult success and the factors that influence cystacanth size within the intermediate host. To assess how host size and intensity of infection influence cystacanth size, infected amphipods (Hyalella azteca) were measured, and sex, length, and width of cystacanths were determined. After a subset of cystacanths was measured, small- and large-size classes of cystacanths were designated. To determine how cystacanth size relates to adult size, green sunfish (Lepomis cyanellus) were fed 10 large or small cystacanths. Fish were dissected 6 wk after infection, and worms were removed. After adult worms were permanently mounted on slides, their length and width were measured. Intensity of infection and amphipod size significantly influenced cystacanth size in that large amphipods harbored larger cystacanths than did small amphipods and heavy infections produced smaller cystacanths than did light infections. Adult worms from the small and large cystacanth-size classes showed no significant difference in size; however, large cystacanths had a significantly higher establishment and survival than did small cystacanths: 40% of large worms and 14% of small worms were recovered. The results of this study indicate that host size and host sharing influence cystacanth size and that cystacanth size is an important factor in determining adult success.     

Virulence of Corynosoma constrictum (Acanthocephala: Polymorphidae) in Hyalella azteca (Amphipoda) throughout parasite ontogeny

Development and growth of parasites depend on resources provided by the host and the parasite's ability to use them. Identifying specific costs incurred by the host provides insight for assessment of parasite energy budgets, which differ among taxa and ontogenetic stages. Data from this study were analyzed using an accelerated failure-time model with intensity as a covariate. Results indicated significantly reduced survival of amphipods, Hyalella azteca, infected with the acanthocephalan Corynosoma constrictum compared with uninfected controls. Male and female amphipod survivorship and infection intensity did not differ; however, amphipods with high-intensity infections (> 16 larvae) died earlier compared with amphipods with low-intensity infections (< 3 larvae). The majority of infected amphipods died between 12 and 24 days postexposure, a period of rapid larval development. It is hypothesized that host death may be due either to an increase in overall larval nutritional demands or to parasite-mediated depletion of a specific host substance. Results from this study suggest that developing C. constrictum satisfies energy requirements by depriving amphipod hosts of resources normally used for somatic growth and maintenance.     

Lack of transmission of Hematodinium sp. in the blue crab Callinectes sapidus through cannibalism

Hematodinium spp. are parasitic dinoflagellates of marine crustaceans. Outbreaks of Hematodinium sp. have impacted commercial landings of the blue crab Callinectes sapidus in the coastal bays of Virginia and Maryland (USA), with seasonal peaks in prevalence reaching 85%. The life cycle and transmission routes of the parasite in blue crabs are poorly understood. Cannibalism and waterborne transmission may be routes of transmission, although little conclusive evidence has been reported for these modes. We examined cannibalism as a route by a series of experiments wherein we repeatedly fed adult and juvenile crabs the tissues of crabs infected with Hematodinium. In each experiment, feeding was done 3 times over the course of 1 wk. Only 2 of 120 crabs were infected within 7 to 9 d after feeding, and these 2 were likely infected prior to the experimental exposures. Crabs inoculated with hemolymph from infected donors served as positive controls. They developed infections over 11 to 21 d, indicating that the Hematodinium sp. used in the cannibalism trials was infectious at the time of inoculation. Because amphipods also harbor Hematodinium-like infections, we fed tissues of infected crabs to the estuarine amphipod Leptocheirus plumulosus. Hematodinium DNA was detected in amphipods shortly after feeding, but not in animals held for longer periods, nor was it observed in histological preparations. Amphipods did not obtain infections by scavenging infected crab tissues. Our results show that Hematodinium sp. is not effectively transmitted through ingestion of diseased tissues, indicating that cannibalism may not be a major route of transmission for Hematodinium sp. in blue crabs.     

Diversity of trematode genetic clones within amphipods and the timing of same-clone infections

The genetic diversity of trematodes within second intermediate hosts has important implications for the evolution of trematode populations as these hosts are utilized after the parasites reproduce asexually within first intermediate hosts and before sexual reproduction within definitive hosts. We characterised the genetic clonal diversity of the marine trematode Maritrema novaezealandensis within amphipod (Paracalliope novizealandiae) second intermediate hosts using four to six microsatellite loci to determine if multiple copies of identical trematode clones existed within naturally infected amphipods. To determine the relative timing of infections by identical clones within hosts, trematode metacercariae were assigned to six developmental stages and the stages of identical clones were compared. The genotypes of 306 trematodes were determined from 44 amphipods each containing more than one trematode. Six pairs of identical trematode clones were recovered in total (representing five amphipods: 11% of amphipods with greater than one trematode) and all pairs of clones belonged to the same developmental stage. This suggests that identical clone infections are effectively synchronous. A general decrease in the number of metacercariae recovered, prevalence, and mean intensity of infection for each subsequent developmental stage coupled with large numbers of metacercariae (>9) only being recovered from recent infections, supports the occurrence of post-infection amphipod mortality and/or within-host trematode mortality. Taken together, our results indicate that natural infections are characterised by high genetic diversity, but that amphipods also periodically encounter "batches" of genetically identical clones, potentially setting the stage for interactions within and between clonal groups inside the host.     

INTERTIDAL AMPHIPODS AS POTENTIAL DISPERSAL AGENTS OF CARPOSPORES OF IRIDAEA LAMINARIOIDES (GIGARTINALES,RHODOPHYTA)1

Mid and late successional benthic algae have poor dispersal capacities. Mobile herbivores may increase dispersal of some algae because spores can survive digestion by grazers and stick to external body appendages. We show that carpospores are the only type of reproductive unit of the rhodophyte Iridaea laminarioides Bory that survive passage through the digestive tract of the amphipod Hyale sp. The spores also stick to the amphipods' legs and are thus carried by the amphipods in the field. Amphipods significantly increased (P < 0.01) the number of spores settling on artificial substrata in places where barriers prevented the normal ingress of algal propagules. The presence of artificial substrata that act as a refuge for the grazers also caused an increase (P < 0.01) in the number of settled spores. Amphipods also significantly increased (P < 0.01) the number of spores under an Ulva sp. canopy in laboratory experiments.     

Parasite-induced suppression of aggregation under predation risk in a freshwater amphipod: Sociality of infected amphipods

Recent findings suggest that grouping with conspecifics is part of the behavioural defences developed by amphipod crustaceans to face predation risk by fish. Amphipods commonly serve as intermediate hosts for trophically transmitted parasites. These parasites are known for their ability to alter intermediate host phenotype in a way that promotes predation by definitive hosts, where they reproduce. If aggregation in amphipods dilutes the risk to be preyed on by fish, then it may dilute the probability of transmission for the parasite using fish as definitive hosts. Using experimental infections, we tested whether infection with the fish acanthocephalan Pomphorhynchus laevis alters attraction to conspecifics in the amphipod intermediate host Gammarus pulex. We also measured G. pulex's activity and reaction to light to detect potential links between changes in aggregation and changes in other behaviours. The attraction to conspecifics in the presence of predator cue, a behaviour found in uninfected gammarids, was cancelled by the infection, while phototaxis was reversed and activity unchanged. We found no correlation between the three behaviours in infected amphipods, while activity and aggregation were negatively correlated in uninfected individuals after the detection of predation cue. The physiological causes and the adaptive value of aggregation suppression are discussed in the context of a multidimensional manipulation.     

Impact of acanthocephalan parasites on aggregation behavior of amphipods (Gammarus pseudolimnaeus)

Acanthocephalan parasites can manipulate the behavior of their amphipod intermediate hosts in ways that increase the amphipod's risk of being eaten by a predator that serves as the final host for the parasite. Some asocial amphipod species have been shown to increase the likelihood of aggregation in response to chemical cues associated with predators. If such aggregation has anti-predation benefits, it might be subject to manipulation by parasites. We tested this hypothesis by comparing the preference of parasitized and unparasitized amphipods (Gammarus pseudolimnaeus) for associating with a group of unparasitized conspecifics, both in the presence and absence of chemical cues from predatory brook sticklebacks (Culaea inconstans). Amphipods with encysted parasites (Corynosoma sp.) avoided aggregating, whereas unparasitized amphipods preferred to aggregate. We also found that the risk of predation by sticklebacks faced by an individual amphipod was significantly lower when the amphipod was in a group compared to when it was alone. This suggests that the aggregation response of unparasitized amphipods is an adaptive response to escape predation. This study provides evidence for a novel parasitic manipulation of intermediate host behavior that is likely to increase transmission to the definitive host.     

The behavioral response of amphipods harboring Corynosoma constrictum (Acanthocephala) to various components of light

Many studies have shown that photic behavior of amphipods is subject to parasitic manipulation. However, all these investigations have focused on but one property of light (i.e., intensity). This study investigated the possibility that variable wavelength sensitivity, as a potentially important component of amphipod ecology, is subject to parasitic manipulation. The photic behavior of freshwater amphipods Hyalella azteca, infected with the duck acanthocephalan Corynosoma constrictum, was tested. The phototactic responses of infected and uninfected amphipods to various wavelengths in the visible spectrum were compared, and to delineate the effects of intensity and wavelength on behavior, the preferences of amphipods for environments characterized by various combinations of light intensity and wavelength were determined. Response to blue light (400-450 nm) was little affected by infection. Amphipod response to higher red region wavelengths (600-700 nm) was altered by infection. Infected amphipods were significantly less responsive to green region light (500-550 nm), which could lead to increased wandering throughout the water column, thereby facilitating increased parasite transmission through increased predation risk. This study reinforces the subtlety with which parasites can alter their host's behavior, presumably resulting in an increased probability of being transmitted from the intermediate host to a definitive host.     

The Influence of Chemical Stimuli from Predators on Precopulatory Pairing by the Amphipod,Gammarus pseudolimnaeus

Many amphipod crustaceans exhibit precopulatory mate guarding. Field samples of the amphipod Gammarus pseudolimnaeus indicated that pairs were positively size assortative. Receptive individuals readily formed pairs in the laboratory and the latency to formation of precopulatory pairs was decreased under threat of predation. In addition, females and, under conditions of extreme danger, males that formed pairs were significantly smaller when under the threat of predation. Amphipods distinguished between chemical stimuli (aquarium water) from predatory and nonpredatory fishes and between chemical cues from fish predators (trout) that had recently eaten conspecific amphipods vs. those fed a control diet of pelleted commercial fish food. These data indicate that chemical stimuli associated with predators can influence reproductive behavior of amphipods. The results also suggest the hypotheses that: 1. search time may be costly in terms of probability of predation; and 2. small pairs may be safer from predation than larger pairs.     

Global diversity of amphipods (Amphipoda; Crustacea) in freshwater

Amphipods are brooding peracaridan crustaceans whose young undergo direct development, with no independent larval dispersal stage. Most species are epibenthic, benthic, or subterranean. There are some 1,870 amphipod species and subspecies recognized from fresh or inland waters worldwide at the end of 2005. This accounts for 20% of the total known amphipod diversity. The actual diversity may still be several-fold. Amphipods are most abundant in cool and temperate environments; they are particularly diversified in subterranean environments and in running waters (fragmented habitats), and in temperate ancient lakes, but are notably rare in the tropics. Of the described freshwater taxa 70% are Palearctic, 13% Nearctic, 7% Neotropical, 6% Australasian and 3% Afrotropical. Approximately 45% of the taxa are subterranean; subterranean diversity is highest in the karst landscapes of Central and Southern Europe (e.g., Niphargidae), North America (Crangonyctidae), and Australia (Paramelitidae). The majority of Palearctic epigean amphipods are in the superfamily Gammaroidea, whereas talitroid amphipods (Hyalella) account for all Neotropic and much of the Nearctic epigean fauna. Major concentrations of endemic species diversity occur in Southern Europe, Lake Baikal, the Ponto-Caspian basin, Southern Australia (including Tasmania), and the south-eastern USA. Endemic family diversity is similarly centered in the Western Palearctic and Lake Baikal. Freshwater amphipods are greatly polyphyletic, continental invasions have taken place repeatedly in different time frames and regions of the world. In the recent decades, human mediated invasions of Ponto-Caspian amphipods have had great impacts on European fluvial ecosystems. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Contrasting patterns of amphipod dispersion in a seagrass meadow between day and night: consistency through a lunar cycle

Changes in light intensity, typically over the course of a day, affect the dispersion of aquatic organisms at short temporal scales. Amphipods, for example, have strong behavioural responses to light conditions. In this study, we used amphipod assemblages inhabiting a Cymodocea nodosa seagrass meadow on the east coast of Gran Canaria Island (eastern Atlantic) to test whether short-term dispersion of seagrass-associated amphipods differed between day and night, testing the consistency throughout an entire lunar cycle. Replicated artificial seagrass units were deployed, and subsequently retrieved, during the day (from 8:00 am to 18:00 pm) and the night (from 18:00 pm to 8:00 am) on three consecutive days within each of the four moon phases of a complete lunar cycle. We collected 13,467 amphipods corresponding to 32 species and 17 families. Significantly larger abundances of amphipods were collected during the night through the entire moon cycle. The total abundance of amphipods was also affected by the moon phases; under full moon, larger abundances of amphipods dispersed into the artificial seagrass units followed by the third quarter, the new moon and the first quarter. The species density of amphipods per unit followed the same pattern. In conclusion, the short-term dispersion of amphipods living in a seagrass meadow was considerably greater during the night than the day, while dispersion of amphipods was more intense under full moon.     

Effects of two acanthocephalan species on the reproduction of Hyalella patagonica (Amphipoda, Hyalellidae) in an Andean Patagonian Lake (Argentina)

The aim of the present study was to evaluate alterations in the reproduction induced by acanthellae and cystacanths of the acanthocephalans Acanthocephalus tumescens and Corynosoma sp. in the amphipod Hyalella patagonica from Lake Mascardi. Specimens of H. patagonica were separated in two categories: paired amphipods (joined specimens during precopulatory mate guarding period until fertilization) and unpaired amphipods (alone specimens). Different analyses were performed: first with paired (n = 406) and unpaired (n = 375) amphipods, and second only with female amphipods (n = 1949), that were classified into three categories (without internal oocytes and eggs, only with internal oocytes, and with eggs). Also, carotenoid extraction was performed of amphipods uninfected (n = 75) and infected (n = 105) by cystacanths of Corynosoma sp. Unpaired amphipods had significantly higher prevalence of cystacanths of both acanthocephalan species than paired ones; but such differences were not found in prevalence of acanthellae. Female amphipods without internal oocytes and eggs showed significantly higher prevalence of cystacanths of both acanthocephalan species than the two other female categories; while females with eggs had significantly higher prevalence of A. tumescens acanthellae. Amphipods infected by Corynosoma sp. showed lower carotenoid concentration than uninfected ones. In Lake Mascardi, there is indirect evidence of both reduced mating success and female fecundity of H. patagonica provoked by both cystacanths species (A. tumescens and Corynosoma sp.). However, infections by acanthellae seem to have no effects.     

About the finding of Pallasiola quadrispinosa (G.O. Sars, 1867) (Gammaridae: Amphipoda) in Lake Vishtynetskoye (Kaliningrad region)

Lake Vishtynetskoye is the only water body in Kaliningrad region where the glacial freshwater relict Pallasiola quadrispinosa (G.O. Sars, 1867) inhabits. Amphipods were recorded in the littoral zone of the lake in autumn and winter 2008. In the biotope with silty shell limestone, amphipods were recorded only in winter (20 ind./m²). The abundance of P. quadrispinosa was 200 ind/m² in autumn and 60 ind/m² in the biotope of sand with overgrowths of water horsetail in winter. At present amphipod, P. quadrispinosa is registered in 12 lakes of Vishtynetsky (Suwalki) Upland.     

Occurrence of Perkinsus sp. in undulated surf clams Paphia undulata from the Gulf of Thailand

The undulated surf clam Paphia undulata supports Thailand's largest shellfishery in the Gulf of Thailand, with landings in 1999 recorded at 70000 t (metric tonnes) yr(-1). We report, for the first time, the prevalence of Perkinsus sp. in clams in the Gulf. A monthly survey from January to December 2001 utilizing the fluid thioglycollate medium (FTM) method showed that average monthly prevalence was 84.7% (n = 360). The monthly percentage of infected clams was generally 100%, with low prevalence in May (66.7%) and no infection in September. The monthly mean infection intensity in terms of Perkinsus sp. cells g(-1) tissue varied from 0 in September to 187 759 +/- 18970 (x +/- SE) in October. No obvious annual variation in intensity and prevalence was observed. Prezoosporangia that developed in FTM were 25 to 75 pm in diameter. A few days after incubation in aerated seawater, the prezoosporangia underwent successive binary cell division and formed motile zoospores (2 to 5 microm long). The zoospores were released into the seawater through a discharge tube formed during the 2- and 4-cell stages. Serial semi-thin sections (1 to 4 pm thickness) of clam tissue (n = 120 clams) showed developing trophozoites 3 to 6 pm in diameter within gills, connective tissue, gonads and, especially, the digestive glands. Microscopic features of different life stages indicated that Perkinsus sp. in Thailand closely resembled P. olseni (= P. atlanticus) reported in Australia, New Zealand, Korea, Japan, Spain and Portugal.     

The distribution and production of Gammarus pseudolimnaeus (Crustacea: Amphipoda) along a reach of the Credit River, Ontario

SUMMARY. The population density of the univoltine amphipod Gammarus pseudolimnaeus was estimated monthly from April 1977 to June 1978 along a 600-m reach using a form of removal sampling at 0.5, 1.5 and 2.5 m from the bank. The probability of catching an amphipod with three sweeps over an area of 0.15 m² was 0.95. Drift and upstream migration were measured for 24 h, also monthly. Population density varied from 1600 m⁻² (newly-hatched young in July) to 26 m⁻² (adults) within 3 m of the bank with 95% confidence limits of 40–60% of the mean; density was highest < 1 m from the bank and beyond 3 m it declined to insignificant levels. No significant difference could be shown between the total catches in the drift and upstream nets over a year and there appeared to be no net migration of amphipods. Mean drift was 2 amphipods day⁻¹.
Growth was estimated from the length composition of monthly samples. Annual production ( P ) was 29.4 kg ha⁻¹ for one generation of the amphipod with a P/B ratio of 4.65. Egg production accounted for only 2% of this estimate. Recruitment was calculated from regressions relating female length to clutch size and egg development time to temperature in the laboratory; mean temperature was measured in the field.
Mortality was greatest in July at the beginning of the generation. It was high again in September and during winter; at these later times it was suggested that starvation occurred in addition to predation. Annual consumption ( C ) of organic matter for the generation calculated from published data on feeding rate was 1547 kg ha⁻¹ which results in a P/C ratio of 1.9%.     

Seasonal and ontogenetic dynamics in trophic transmission of parasites

Transmission rates from the intermediate (amphipods) to the definitive hosts (fish) were quantified for two helminth species ( Cyathocephalus truncatus , Cestoda, and Cystidicola farionis , Nematoda) both seasonally and through the ontogeny of the final hosts (arctic charr, Salvelinus alpinus , and brown trout, Salmo trutta ). Amphipods ( Gammarus lacustris ) were important prey for both fish species, especially in the autumn. Both parasite species had low infection levels in amphipods compared to high abundance in fish. The seasonal variations in transmission rate of C. truncatus procercoids from amphipods to fish were in accordance with the observed abundance in fish hosts, being highest in the autumn and lowest during late winter and early summer. During summer, however, the estimated monthly transmission rates of C. truncatus were higher than the observed infection levels in the fish, suggesting restricted establishment success and shorter development time and longevity of the cestode in fish at higher temperatures. The accumulated transmission of C. farionis over the ontogeny of arctic charr was similar to the observed age-specific infection levels, reflecting a high establishment success and longevity of this parasite in charr. In contrast, brown trout exhibited an infection level that was much lower than the estimated transmission rates, suggesting a high resistance against C. farionis in these fish. The magnitudes of the estimated transmission rates were sufficient to explain the paradoxical contrast between low infection levels in the intermediate and high in the final hosts.