REGULAR PAPERS Unusual stable isotope fractionation patterns observed for fish host—parasite trophic relationships

Trophic relationships between four taxa of fish parasite (Cestoda, Nematoda, Isopoda, Copepoda) and their hosts ( Gasterosteus aculeatus, Merlangius merlangus, Boops boops, Platichthys flesus ) were investigated using stable isotopes of carbon and nitrogen. δ¹⁵N differences between parasite and host were unlike those conventionally observed among consumers and their diets. In no case were parasites enriched in 15N with respect to the host organism; endoparasites were significantly and consistently depleted. In no case were parasites enriched in 15N with respect to the host organism; endoparasites were significantly and consistently depleted.     

Parasite-induced changes in nitrogen isotope signatures of host tissues

To estimate isotopic changes caused by trematode parasites within a host, we investigated changes in the carbon and nitrogen isotope ratios of the freshwater snail Lymnaea stagnalis infected by trematode larvae. We measured carbon and nitrogen stable isotopes within the foot, gonad, and hepatopancreas of both infected and uninfected snails. There was no significant difference in the delta13C and delta15N values of foot and gonad between infected and uninfected snails; thus, trematode parasite infections may not cause changes in snail diets. However, in the hepatopancreas, delta15N values were significantly higher in infected than in uninfected snails. The 15N enrichment in the hepatopancreas of infected snails is caused by the higher 15N ratio in parasite tissues. Using an isotope-mixing model, we roughly estimated that the parasites in the hepatopancreas represented from 0.8 to 3.4% of the total snail biomass, including the shell.     

Nested assemblages resulting from host size variation: the case of endoparasite communities in fish hosts

Nested species subsets are a common pattern in many types of communities found in insular or fragmented habitats. Nestedness occurs in some communities of ectoparasites of fish, as does the exact opposite departure from random assembly, anti-nestedness. Here, we looked for nested and anti-nested patterns in the species composition of communities of internal parasites of 23 fish populations from two localities in Finland. We also compared various community parameters of nested and anti-nested assemblages of parasites, and determined whether nestedness may result simply from a size-related accumulation of parasite species by feeding fish hosts. Nested parasite communities were characterised by higher prevalence (proportion of infected fish) and intensities of infection (number of parasites per fish) than anti-nested communities; the two types of non-random communities did not differ with respect to parasite species richness, however. In addition, the correlation between fish size and the number of parasite species harboured by individual fish was much stronger in nested assemblages than in anti-nested ones, where it was often nil. These results were shown not to be artefacts of sampling effort or host phylogeny. They apply to both assemblages of adult and larval parasites, which were treated separately. Since species of larval parasites are extremely unlikely to interact with one another in fish hosts, the establishment of nestedness appears independent of the potential action of interspecific interactions. The species composition of these parasite communities is not determined from within the community, but rather by the extrinsic influence of host feeding rates and how they amplify differences among parasite species in probabilities of colonisation or extinction. Nested patterns occur in parasite communities whose fish hosts accumulate parasites in a predictable fashion proportional to their size, whereas anti-nested communities occur in parasite communities whose fish hosts do not, possibly because of dietary specialisation preventing them from sampling the entire pool of parasite species available locally. Thus, nestedness in parasite communities may result from processes somewhat different from those generating nested patterns in free-living communities.     

Assessing nursery habitat quality for native smelts (Osmeridae) in the low-salinity zone of the San Francisco estuary

Habitat quality was assessed for two native osmerids, delta smelt Hypomesus transpacificus and longfin smelt Spirinchus thaleichthys , between two distinct nursery areas located in the low-salinity zone of the San Francisco estuary. The relationship between several variables was investigated including fish density, fish size, feeding success and the general condition of larvae as well as juveniles for both species. The nursery habitats that were evaluated included the North and South Channels of Suisun Bay. The results showed higher densities of zooplankton and decreased water velocities for the North Channel when compared to the South Channel. The dominant prey item was calanoid copepod Pseudodiaptomus forbesi for both species although longfin smelt residing in the North Channel also included another copepod in their diets, Acanthocyclops spp. In both locations, delta smelt fed predominantly during daytime flood tides, while longfin smelt feeding appeared to continue into the night hours. When both locations were compared, delta smelt in the North Channel exhibited higher densities, larger sizes, increased somatic condition and larvae <15 mm standard length demonstrated greater feeding success. Longfin smelt, exhibited similar densities, feeding success and size distributions between both channels, but generally showed poorer somatic condition for the South Channel, potentially due to energetic costs associated with documented vertical migration behaviour. Overall, the physical conditions of the North Channel provided superior habitat for both species, while the South Channel afforded only marginal habitat for longfin smelt and very poor habitat for delta smelt. Therefore, the North Channel of Suisun Bay acts as critical nursery habitat by providing better feeding and growing conditions leading to increased health and survival for both species.     

Comparison of small subunit ribosomal RNA gene and internal transcribed spacer sequences among isolates of the intranuclear microsporidian Nucleospora salmonis

Nucleospora salmonis is an intranuclear microsporidian associated with a proliferative disorder of the lymphoid cells of captive salmonid fish in the northwestern and northeastern regions of North America, in France, and in Chile. Newer diagnostic approaches have used the polymerase chain reaction (PCR) to detect the parasite in fish tissues. The target sequences for these assays lie in the small subunit ribosomal RNA (ssu rRNA) gene or internal transcribed spacer (ITS) as determined from N. salmonis from chinook salmon (Oncorhynchus tshawytscha) from the Pacific Northwest of North America. The lack of sequence data on parasites from diverse geographic origins and hosts led us to compare several isolates of N. salmonis. There was a high degree of similarity in the ssu rDNA sequences (> 98%) among all the isolates of N. salmonis examined, regardless of host or geographic origin. The greatest sequence differences were found between isolates from the Pacific regions of America. Isolates from Chile shared sequences with one or both geographic groups from North America. A similar distribution of sequence types was observed when ITS-1 sequences of selected isolates were analyzed. Sequence data from two N. salmonis-like isolates from marine non-salmonid fish showed one closely related and the second less closely related to N. salmonis isolates from salmonid fish. These results provide evidence for a homogeneous group of aquatic members of the genus Nucleospora found among salmonid fish (N. salmonis) that can be detected using diagnostic PCR assays with ssu rDNA target sequences. The presence of parasites related to N. salmonis among marine fish suggests a potentially broad host and geographic distribution of members of the family Enterocytozoonidae.     

Morphology and pathology of the ectoparasitic copepod, Nicothoë astaci ('lobster louse') in the European lobster, Homarus gammarus

Ectoparasitic copepods have been reported in a wide range of aquatic animals, including crustacean shellfish. However, with the exception of the salmon louse, Lepeophtheirus salmonis, our knowledge of such parasites in commercial species is rudimentary. The current study examines the morphology and pathology of the parasitic copepod, Nicotho? astaci (the 'lobster louse') in its host, the European lobster, Homarus gammarus. Lobsters were sampled from waters surrounding Lundy Island (Bristol Channel, UK) and all individuals collected were found to harbour female adult N. astaci in their gills, with a mean of 47·3 parasites/lobster. The majority of N. astaci were found in the basal region of pleurobranch gills. The parasite was found to attach to gill filaments via its oral sucker, maxillae and maxillipeds, and to feed on host haemolymph (blood) through a funnel-like feeding channel. It caused varying degrees of damage to the host gill, including occlusion of gill filaments and disruption to the vascular system in the central axis. Although there was evidence of extensive host response (haemocytic infiltration) to the parasite, it was displaced from the parasite attachment site and thus was observed in the central gill axis below. The region of gill filament immediately underlying the parasite feeding channel was devoid of such activity suggesting that the parasite interferes with the cellular defence and haemostatic mechanisms of the lobster in order to maintain invasion of the host.     

Are eels (Anguilla anguilla L.) planktonic feeders? Evidence from parasite communities

The metazoan parasite communities of eels Anguilla anguilla were investigated in three riverine localities and one lake in Devon. Altogether 21 species of parasite, the majority of the British eel parasite fauna, were recorded. Parasite communities in the three riverine localities were more similar to each other than to that of the lake. The four parasite component communities were typical and representative of those from eels in other parts of the U.K. They were characterized by low species diversity and domination by a single species: Pseudodacytlogyrus anguillae in the rivers and Diplostomum spathaceum in the lake. Both the dominant species and the majority of parasite species, and hence individuals, in any locality infected eels directly by free swimming planktonic larval stages or indirectly by the ingestion of infected planktonic copepod intermediate hosts. Examination of other species of fish in each locality showed that only Raphidascaris acus utilized a fish species as an intermediate or paratenic host to infect eels. Increases in prevalence and intensity of infection with size of eel by helminths such as Camallanus lacustris and Paraquimperia tenerrima were observed in localities in which transmission of these parasites could not have involved another fish species. Over 63% of the specialist, specific parasites of eels throughout Europe employ planktonic transmission to their hosts, as do several species of the commoner generalists. A study of eel parasite communities thus leads to the conclusion that eels are widely and closely associated in their habits with plankton and that eels of all sizes feed regularly upon planktonic copepods: the widely accepted view of eels as being more or less exclusively benthic in habit and feeding behaviour requires re-appraisal.     

Pathological and ecological host consequences of infection by an introduced fish parasite

The infection consequences of the introduced cestode fish parasite Bothriocephalus acheilognathi were studied in a cohort of wild, young-of-the-year common carp Cyprinus carpio that lacked co-evolution with the parasite. Within the cohort, parasite prevalence was 42% and parasite burdens were up to 12% body weight. Pathological changes within the intestinal tract of parasitized carp included distension of the gut wall, epithelial compression and degeneration, pressure necrosis and varied inflammatory changes. These were most pronounced in regions containing the largest proportion of mature proglottids. Although the body lengths of parasitized and non-parasitized fish were not significantly different, parasitized fish were of lower body condition and reduced weight compared to non-parasitized conspecifics. Stable isotope analysis (δ(15)N and δ(13)C) revealed trophic impacts associated with infection, particularly for δ(15)N where values for parasitized fish were significantly reduced as their parasite burden increased. In a controlled aquarium environment where the fish were fed ad libitum on an identical food source, there was no significant difference in values of δ(15)N and δ(13)C between parasitized and non-parasitized fish. The growth consequences remained, however, with parasitized fish growing significantly slower than non-parasitized fish, with their feeding rate (items s(-1)) also significantly lower. Thus, infection by an introduced parasite had multiple pathological, ecological and trophic impacts on a host with no experience of the parasite.     

Evolutionary implications of the adaptation to different immune systems in a parasite with a complex life cycle

Many diseases are caused by parasites with complex life cycles that involve several hosts. If parasites cope better with only one of the different types of immune systems of their host species, we might expect a trade-off in parasite performance in the different hosts, that likely influences the evolution of virulence. We tested this hypothesis in a naturally co-evolving host-parasite system consisting of the tapeworm Schistocephalus solidus and its intermediate hosts, a copepod, Macrocyclops albidus, and the three-spined stickleback Gasterosteus aculeatus. We did not find a trade-off between infection success in the two hosts. Rather, tapeworms seem to trade-off adaptation towards different parts of their hosts' immune systems. Worm sibships that performed better in the invertebrate host also seem to be able to evade detection by the fish innate defence systems, i.e. induce lower levels of activation of innate immune components. These worm variants were less harmful for the fish host likely due to reduced costs of an activated innate immune system. These findings substantiate the impact of both hosts' immune systems on parasite performance and virulence.     

Metazoan parasites of sticklebacks on Sable Island, Northwest Atlantic Ocean: biogeographic considerations

Three species of sticklebacks ( Apeltes quadracus, Gasterosteus aculeatus , and Pungitius pungitius ( n = 236) were collected from five ponds on Sable Island. The nematodes Pseudoterranova decipiens, Contracaecum sp., Paracuaria adunca , and Cosmucephalus obvelatus , and the cestode Diphyllobothrium ditremum parasitized three-spined sticklebacks ( G. aculeatus ) and four-spined sticklebacks ( A. quadracus ) inhabiting four brackish water ponds. All the parasites except P. decipiens infected nine-spined sticklebacks ( P. pungitius ) from a freshwater pond. In addition, the cestode Schistocephalus pungitii , the copepod Thersitina gasterostei , and the monogenean Gyrodactylus canadensis occurred in nine-spined sticklebacks from the freshwater pond. The two cestodes, the copepod, and the sealworm, P. decipiens , were the most common parasites encountered. The remaining helminths were relatively rare. Most of the parasite species were larval forms which use gulls or seals as definitive hosts. These parasites probably colonized Sable Island with their definitive hosts, whereas only two species ( T. gasterostei and G. canadensis ) successfully colonized the island ponds with their fish hosts. The low parasite species richness encountered is attributed to the impoverished nature of the host fauna of Sable island, and the difficulty of colonization as a result of the island's isolation with respect to the mainland.     

Infracommunity structure of parasites of Hemigymnus melapterus (Pisces: Labridae) from Lizard Island, Australia: the importance of habitat and parasite body size

This study describes the community of all metazoan parasites from 14 individuals of thicklip wrasse, Hemigymnus melapterus, from Lizard Island, Australia. All fish were parasitized, and 4,649 parasite individuals were found. Twenty-six parasite species were identified although only 6 species were abundant and prevalent: gnathiid isopods, the copepod Hatschekia hemigymni, the digenean Callohelmis pichelinae, and 3 morphotypes of tetraphyllidean cestode larvae. We analyzed whether the body size and microhabitat of the parasites and size of the host affected understanding of the structure of the parasite community. We related the abundance, biovolume, and density of parasites with the host body size and analyzed the abundances and volumetric densities of some parasite species within microhabitats. Although the 2 most abundant species comprised 75% of all parasite individuals, 4 species, each in similar proportion, comprised 85% of the total biovolume. Although larger host individuals had higher richness, abundance, and biovolume of parasites than smaller individuals, overall parasite volumetric density actually decreased with the host body size. Moreover, parasites exhibited abundances and densities significantly different among microhabitats; some parasite species depended on the area available, whereas others selected a specific microhabitat. Parasite and habitat size exhibited interesting relationships that should be considered more frequently. Considerations of these parameters improve understanding of parasite community structure and how the parasites use their habitats.     

An attempt to balance the energy budget of a host-parasite system

A quantitative assessment of the demand by the parasite Schistocephalus solidus (Miiller, 1776) upon its secondary host, Gasterosteus aculentus L., was obtained by comparing the energy budgets for infected and non-infected fish. Observations on the energy trans-formations during feeding, assimilation, growth and respiration of fishes indicated some overall effects of the parasite upon host metabolism.
Infection resulted in a greater depletion of host food reserws, shown by a marked in-crease in mortality of parasitized fish during starvation. When fed ad libitum upon Tubifex , differences were recorded in the feeding and assimilation rates of fish. By comparison, no significant differences were detected in the respiratory expenditure of infected and non-infected hosts. The computation of energy budgets indicated that fish bearing parasites characteristically exhibited a higher gross efficiency than did fish without parasites. How-ever, subtraction of the calculated effect due to the presence of worms, suggested that the efficiency of infected fish alone, that is, without their parasites, was actually lower than the efficiency of uninfected fish. It is considered therefore that the apparent greater energy turnover in a parasitized fish is due to the parasite being more efficient in its energy trans-formations than is its host.     

Role of cellular response in elimination of the monogenean Neoheterobothrium hirame in Japanese flounder Paralichthys olivaceus

Adult worms of the blood-feeding monogenean parasite Neoheterobothrium hirame, which cause anemia in the Japanese flounder Paralichthys olivaceus, attach to the host fish by embedding their posterior part deeply into the host tissue. To investigate the possibility that cellular responses of the host fish can eliminate N. hirame, flounder were experimentally infected with N. hirame larvae and reared in either fed or starved conditions. Mature parasites were identified on the buccal cavity wall of the fish 33 d post-infection (Day 33). Monocytes/macrophages and granulocytes increased rapidly in the blood and infected sites after the appearance of mature parasites. These cells adhered to the tegument of the parasites. In addition, a few cells with large electron-dense granules (DGCs) were observed in the inflammatory foci. On Day 47, the tegument of some parasites collapsed partially and were phagocytosed by the infiltrated host cells. Some infiltrated cells adhered directly to the inner tissues of the parasites. On Day 54, in the fed fish group, the loss of the tegument led to damage of the parasites' inner tissue by a large number of infiltrated cells. In this group, the elimination of the parasites was noted from Day 47 to 54. These observations probably suggest that the cellular response of the host fish destructed the parasite's posterior part embedded in the tissue, thereby eliminating the parasites. On the other hand, a high mortality was observed in the starved group. The starved fish developed much more severe anemia than the fed fish, and the elimination of the parasites was not observed in this group. The results of the present study suggest that flounder can eliminate N. hirame if they are fed sufficiently.     

The association between parasite infection and growth rates in Arctic charr: do fast growing fish have more parasites?

Trophically transmitted parasites are known to impair fish growth in experimental studies, but this is not well documented in natural populations. For Arctic charr [Salvelinus alpinus (L.)], individual growth is positively correlated with food consumption. However, increased food consumption will increase the exposure to trophically transmitted parasites. Using a correlative approach, we explore the association between parasite abundance and the individual growth of Arctic charr from five lakes within the same watercourse. The studied parasite species differ in their life cycles and cost to the host. We predicted a positive association between parasite abundance and fish growth for parasites of low pathogenicity reflecting high consumption rates, and a negative association at higher parasite abundances for more costly parasites. We found no direct negative associations between parasite abundance and fish growth. The relationship between parasite abundance and growth was linearly positive for the low costly Crepidostomum sp. and concave for the more costly Eubothrium salvelini. In natural fish populations, the negative effects of parasites on fish growth might be outweighed by the energy assimilated from feeding on the intermediate host. However, experimental studies with varying food consumption regimes are needed to determine the mechanisms underlying our observations.

     

Microhabitat distributions and species interactions of ectoparasites on the gills of cichlid fish in Lake Victoria,Tanzania

Heterogeneous exposure to parasites may contribute to host species differentiation. Hosts often harbour multiple parasite species which may interact and thus modify each other’s effects on host fitness. Antagonistic or synergistic interactions between parasites may be detectable as niche segregation within hosts. Consequently, the within-host distribution of different parasite taxa may constitute an important axis of infection variation among host populations and species. We investigated the microhabitat distributions and species interactions of gill parasites (four genera) infecting 14 sympatric cichlid species in Lake Victoria, Tanzania. We found that the two most abundant ectoparasite genera (the monogenean Cichlidogyrus spp. and the copepod Lamproglena monodi) were non-randomly distributed across the host gills and their spatial distribution differed between host species. This may indicate microhabitat selection by the parasites and cryptic differences in the host–parasite interaction among host species. Relationships among ectoparasite genera were synergistic: the abundances of Cichlidogyrus spp. and the copepods L. monodi and Ergasilus lamellifer tended to be positively correlated. In contrast, relationships among morphospecies of Cichlidogyrus were antagonistic: the abundances of morphospecies were negatively correlated. Together with niche overlap, this suggests competition among morphospecies of Cichlidogyrus. We also assessed the reproductive activity of the copepod species (the proportion of individuals carrying egg clutches), as it may be affected by the presence of other parasites and provide another indicator of the species specificity of the host–parasite relationship. Copepod reproductive activity did not differ between host species and was not associated with the presence or abundance of other parasites, suggesting that these are generalist parasites, thriving in all cichlid species examined from Lake Victoria.     

Isotopic discrimination between food and blood and feathers of captive penguins: implications for dietary studies in the wild

Using measurements of naturally occurring stable isotopes to reconstruct diets or source of feeding requires quantifying isotopic discrimination factors or the relationships between isotope ratios in food and in consumer tissues. Diet-tissue discrimination factors of carbon ((13)C/(12)C, or delta (13)C) and nitrogen ((15)N/(14)N, or delta (15)N) isotopes in whole blood and feathers, representing noninvasive sampling techniques, were examined using three species of captive penguins (king Aptenodytes patagonicus, gentoo Pygoscelis papua, and rockhopper Eudyptes chrysocome penguins) fed known diets. King and rockhopper penguins raised on a constant diet of herring and capelin, respectively, had tissues enriched in (15)N compared to fish, with discrimination factors being higher in feathers than in blood. These data, together with previous works, allowed us to calculate average discrimination factors for (15)N between whole lipid-free prey and blood and feathers of piscivorous birds; they amount to +2.7 per thousand and +4.2 per thousand, respectively. Both fish species were segregated by their delta (13)C and delta (15)N values, and importantly, lipid-free fish muscle tissue was consistently depleted in (13)C and enriched in (15)N compared to whole lipid-free fish. This finding has important implications because previous studies usually base dietary reconstructions on muscle of prey rather than on whole prey items consumed by the predator. We tested the effect of these differences using mass balance calculations to the quantification of food sources of gentoo penguins that had a mixed diet. Modeling indicated correct estimates when using the isotopic signature of whole fish (muscle) and the discrimination factors between whole fish (muscle) and penguin blood. Conversely, the use of isotopic signatures of muscle together with discrimination factors between whole fish and blood (or the reverse) leads to spurious estimates in food proportions. Consequently, great care must be taken in the choice of isotopic discrimination factors to apply to wild species for which no controlled experiments on captive individuals have been done. Finally, our results also indicate that there is no need to remove lipids before isotopic analysis of avian blood.     

Living off a fish: a trade-off between parasites and the immune system

Research in fish immune system and parasite invasion mechanisms has advanced the knowledge of the mechanisms whereby parasites evade or cope with fish immune response. The main mechanisms of immune evasion employed by fish parasites are reviewed and considered under ten headings. 1) Parasite isolation: parasites develop in immuno-privileged host tissues, such as brain, gonads, or eyes, where host barriers prevent or limit the immune response. 2) Host isolation: the host cellular immune response isolates and encapsulates the parasites in a dormant stage without killing them. 3) Intracellular disguise: typical of intracellular microsporidians, coccidians and some myxosporeans. 4) Parasite migration, behavioural and environmental strategies: parasites migrate to host sites the immune response has not yet reached or where it is not strong enough to kill them, or they accommodate their life cycles to the season or the age in which the host immune system is down-regulated. 5) Antigen-based strategies such as mimicry or masking, variation and sharing of parasite antigens. 6) Anti-immune mechanisms: these allow parasites to resist innate humoral factors, to neutralize host antibodies or to scavenge reactive oxygen species within macrophages. 7) Immunodepression: parasites either suppress the fish immune systems by reducing the proliferative capacity of lymphocytes or the phagocytic activity of macrophages, or they induce apoptosis of host leucocytes. 8) Immunomodulation: parasites secrete or excrete substances which modulate the secretion of host immune factors, such as cytokines, to their own benefit. 9) Fast development: parasites proliferate faster than the ability of the host to mount a defence response. 10) Exploitation of the host immune reaction. Knowledge of the evasion strategies adopted by parasites will help us to understand host-parasite interactions and may therefore help in the discovery of novel immunotherapeutic agents or targeted vaccines, and permit the selection of host-resistant strains.     

Community ecology of metazoan parasites of the anchovy Anchoa tricolor (Osteichthyes: Engraulidae) from the coastal zone of the State of Rio de Janeiro, Brazil.

Between October 2001 and March 2002, 103 specimens of A. tricolor from Angra dos Reis (23 degrees 01' S, 44 degrees 19' W), in the coastal zone of the State of Rio de Janeiro, Brazil, were analyzed in order to study their metazoan parasite infracommunities. Ten species of metazoan parasites were collected: 4 digeneans, 1 cestode, 1 acantocephalan, 2 nematodes, 1 copepod, and 1 hirudinean; 77.7% of the fishes were parasitized by one or more metazoan, with a mean of 3.5 +/- 6.2 parasite/fish. Digenean was the most dominant with 4 species that accounted for 53.2% of the total parasites collected; Ergasilus sp. was the most abundant species. Abundance and prevalence of Parahemiurus merus (Linton, 1910) were positively correlated with the total length of host. Relationships between total body length of fish and both total parasite abundance and mean parasite species richness were observed. Mean parasite diversity of species was correlated to host's total length, with significant differences found between male and female fishes. Two pairs of larval species showed significant positive association and covariation. The metazoan parasite infracommunities of A. tricolor presented dominance of larval endoparasites; correlation of parasite abundance, diversity, and species richness with host total length; and low number of parasite interspecific relationships. The parasite community of A. tricolor showed some similarities with the parasite community of another South American Atlantic engraulid.     

Fractionation of δ15N and δ13C for Atlantic salmon and its intestinal cestode Eubothrium crassum

Stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) were measured for Atlantic salmon Salmo salar and their intestinal cestode, Eubothrium crassum , sharing the same diet. Atlantic salmon muscle tissues were enriched in ¹⁵N and depleted in ¹³C compared to their prey (sprat Sprattus sprattus sprattus ) and their intestinal cestode. There was no significant difference in δ¹⁵N or δ¹³C between E. crassum and the sprat. Differences in nutrient uptake and intestine physiology between Atlantic salmon and E. crassum are discussed, as well as how these may give rise to different fractionations of stable isotopes between a host and its parasites. Furthermore, Atlantic salmon contained a significantly higher lipid content than their prey, which may partly explain differences in δ¹³C values between the host and its cestode. In addition, cestodes inhabiting lipid-rich hosts were also lipid rich. Larger Atlantic salmon were enriched in ¹⁵N compared to smaller fish. Cestodes inhabiting large hosts were also enriched in ¹⁵N compared to parasites living in smaller hosts. The last two results were explained by larger fish possibly feeding from a higher trophic level, or from larger and older prey, that resulted in both a higher lipid content and an enrichment in ¹⁵N.     

The role of the European bitterling (<Emphasis Type="Italic">Rhodeus amarus,</Emphasis> Cyprinidae) in parasite accumulation and transmission in riverine ecosystems

In aquatic ecosystems, fish play a key role in parasite accumulation and transmission to predacious animals. In the present study, realized on seven populations of a small cyprinid fish species, the European bitterling Rhodeus amarus, we investigated (1) the role of the European bitterling as a potential intermediate or paratenic host, (2) the ability of the fish to accumulate parasites with similar final host group, and (3) its significance as a potential source of parasite infection in the ecosystem in respect to habitat characteristics. A total of 36 parasite species were recorded; 31 species (90% of all parasite specimens) were classified as endoparasites. Most of the endoparasites were found in the larval life stage, using bitterling as an intermediate or paratenic host. In particular, parasite community structure showed significantly higher proportions of allogenic parasites in comparison with autogenic. The supposed co-occurrence of parasite species with identical final host groups showed only a weak association. The adjacent reservoir areas were a significant determinant of both the total and infracommunity parasite species richness and for the mean parasite abundance. No relationship between the distance of sampling site from the adjacent reservoir and parasite community characteristics was found. As a small-sized fish with a wide distribution range and high local abundances, the European bitterling can represent a natural prey for a wide range of piscivorous predators. Due to its susceptibility to the number of larval endoparasites, this fish species may therefore fulfill the role as important transmitter of parasites to their final hosts.