Carbohydrate localization on Gyrodactylus salaris and G. derjavini and corresponding carbohydrate binding capacity of their hosts Salmo salar and S. trutta

The congeners Gyrodactylus salaris and G. derjavini are specific ectoparasites of Atlantic salmon Salmo salar and brown trout S. trutta, respectively. To elucidate the involvement of lectin-carbohydrate interactions in this host specificity, carbohydrates on the tegument of the two species and the corresponding lectin activity of their hosts have been studied. Carbohydrate composition on the tegument differed significantly between the two gyrodactylids. Three of four commercially available peroxidase-labelled lectins with primary affinity towards D-mannoside, D-GalNAc and L-fucose bound more strongly to G. derjavini than to G. salaris. Lectins with an affinity towards D-mannoside and D-GalNAc bound significantly stronger to the cephalic lobes on G. derjavini compared to the tegument and sheaths of the hamuli. One brown trout strain and three different salmon strains were tested for lectin activity in skin and plasma. Two Baltic salmon strains and one strain from the Atlantic region were included. Brown trout differed significantly from the salmon strains when skin samples were tested for D-GalNAc activity. Lectins binding to other carbohydrates showed trends for similar host differences. The implications of carbohydrate-lectin interactions for host specificity in gyrodactylids are discussed.     

The susceptibility of grayling (Thymallus thymallus) to experimental infections with the monogenean Gyrodactylus salaris

The pathogenic monogenean Gyrodactylus salaris infecting Atlantic salmon (Salmo salar) is found to attach and reproduce under laboratory conditions on several species in the subfamily Salmoninae other than the Atlantic salmon. The gyrodactylid species Gyrodactylus thymalli infecting grayling (Thymallus thymallus) in another subfamily, Thymallinae, is previously said to be very similar to G. salaris based on morphometry and genetical analysis which prompted the present laboratory experiments to test the susceptibility and resistance of grayling to G. salaris. All 0+ and 1+ grayling became infected with G. salaris during the experimental infection procedure. However, both innate resistant and susceptible grayling were found. In susceptible individually isolated fish, parasite reproduction lasted for more than 35 days. Parasite reproduction also occurred among grouped grayling as judged from the duration of infection of more than 50 days. However, grayling susceptibility as judged from G. salaris reproduction, was very limited. Hence, the results indicate significant biological differences between the function of Atlantic salmon and grayling as host for G. salaris. The grayling is interpreted as unable to sustain G. salaris in nature which implies that G. thymalli is not conspecific with G. salaris. However, G. salaris dispersal by grayling cannot be excluded.     

Qualitative analysis of the risk of introducing Gyrodactylus salaris into the United Kingdom

Gyrodactylus salaris is a freshwater, monogenean ectoparasite of Baltic strains of Atlantic salmon Salmo salar on which it generally causes no clinical disease. Infection of other strains of Atlantic salmon in Norway has resulted in high levels of juvenile salmon mortality and highly significant reductions in the population. The parasite is a major exotic disease threat to wild Atlantic salmon in the UK. This paper qualitatively assesses the risk of introduction and establishment of G. salaris into the UK. The current UK fish health regime prevents the importation of live salmonids from freshwater in territories that have not substantiated freedom from G. salaris. The importation of other species, e.g. eels Anguilla anguilla and non-salmonid fish, represents a low risk because the likelihood of infection is very low and the parasite can only survive on these hosts for less than 50 d. Importation of salmon carcasses presents a negligible risk because harvested fish originate from seawater sites and the parasite cannot survive full strength salinity. The importation of rainbow trout Oncorhynchus mykiss carcasses from G. salaris infected freshwater sites might introduce the parasite, but establishment is only likely if carcasses are processed on a salmonid farm in the UK. A number of mechanical transmission routes were considered (e.g. angling equipment, canoes, ballast water) and the most important was judged to be the movement of live fish transporters from farms on mainland Europe direct to UK fish farms. In the future, territories may have to substantiate freedom from G. salaris and economic drivers for live salmonid imports may strengthen. Under these circumstances, legal or illegal live salmonid imports would become the most significant risk of introduction.     

Mitochondrial DNA variation of Gyrodactylus spp (Monogenea, Gyrodactylidae) populations infecting Atlantic salmon, grayling, and rainbow trout in Norway and Sweden

Approximately 800 bp of the mitochondrial cytochrome oxidase I (COI) gene were sequenced from 76 Gyrodactylus specimens of 32 salmonid host populations, i.e. from Salmo salar, Thymallus thymallus, and Oncorhynchus mykiss in Norway, Sweden and Latvia. The COI sequences indicated a substantial intraspecific differentiation of Gyrodactylus salaris and Gyrodactylus thymalli. In total, 12 haplotypes were identified which group into five well supported clades, three clades with parasites from Atlantic salmon and two clades with parasites from grayling. The basal nodes linking the five clades together are only weakly supported. Thus, there is no support for the monophyly of all G. salaris haplotypes and the monophyly of all G. thymalli haplotypes. The lack of monophyly of the mitochondrial haplotypes of G. salaris and G. thymalli may indicate that G. salaris and G. thymalli represent (i). two polytypic species or (ii). one polytypic species, or (iii). refer to a complex of more than two sibling species. The mtDNA data indicate multiple introductions of G. salaris and G. thymalli into Norway. A minimum of three independent introductions of G. salaris and two independent introductions of G. thymalli are supported. This is congruent with earlier hypotheses on the introduction of G. salaris and G. thymalli into Norway.     

Demonstrating freedom from Gyrodactylus salaris (Monogenea: Gyrodactylidae) in farmed rainbow trout Oncorhynchus mykiss

This paper describes an approach to demonstrate freedom of individual rainbow trout farms from Gyrodactylus salaris Malmberg, 1957. The infection status of individual farms is relevant should G. salaris be introduced into a country or zone previously known to be free of the parasite. Trade from farms where G. salaris may have been introduced would be restricted until freedom had been demonstrated. Cage, fish and parasite sample sizes were calculated based on the minimum detectable prevalence (P*), test characteristics, population size, and Type I and II errors. Between 5 and 23 cages per farm would need to be sampled to demonstrate freedom at a cage level P* of 10%. The number of fish sampled per cage depended mainly on the test sensitivity (probability of correctly identifying an infected fish). Assuming a test sensitivity of 99% at the fish level, 59 fish per cage are needed (P* = 5%). Since G. salaris may exist in mixed infection with G. derjavini, testing a sample of gyrodactylid parasites may not result in the parasite being detected when present. Test sensitivity at the fish level depends on the number of gyrodactylids on the fish, the proportion of which are G. salaris and the number examined. Assuming a P* of 5% (i.e. G. salaris are at least 5% of the gyrodactylid population), between 20 and 73 parasites per fish would need to be sampled (depending on abundance) to maintain the Type I error at 0.01 (thus a fish level test sensitivity of 99%). This work identifies the critical information, and further research, needed to assess freedom from G. salaris with a known level of confidence; this is essential to provide a sound scientific basis for decision-making about disease control measures.     

Initial steps of speciation by geographic isolation and host switch in salmonid pathogen Gyrodactylus salaris (Monogenea: Gyrodactylidae)

To test the hypothesis that host-switching can be an important step in the speciation of gyrodactylid monogenean flatworms, we inferred the phylogeny within a cluster of parasites morphologically close to Gyrodactylus salaris Malmberg 1957, collected from Atlantic, Baltic and White Sea salmon (Salmo salar), farmed rainbow trout (Oncorhynchus mykiss), and grayling (Thymallus thymallus) from Northern Europe. The internal transcribed spacer region of the nuclear ribosomal gene was sequenced for taxonomic identification. Parasites on grayling from the White Sea Basin differed from the others by one nucleotide (0.08%), the remainder were identical to the sequence published earlier from Norway (G. salaris on salmon), England (Gyrodactylus thymalli on grayling), and the Czech Republic (unidentified salaris/thymalli on trout). For increased resolution, 813 nucleotides of the mitochondrial COI gene of 88 parasites were sequenced and compared with 76 published sequences using phylogenetic analysis. For all tree building algorithms (NJ, MP), the parasites formed a star-like phylogeny of six definite sister clades, indicating nearly simultaneous radiation. Average K2P distances between clades were 1.8-2.6%, and internal mean distances 0.2-1.1%. The genetic distance to the nearest known relative, Gyrodactylus lavareti Malmberg, was 24%. A variable salmon-specific mitochondrial Clade I was observed both in the Baltic Basin and in pathogenic populations introduced to the Atlantic and White Sea coasts. An invariable Clade II was common in rainbow trout farms in Sweden, Denmark and Finland; the same haplotype was also infecting salmon in a landlocked population in Russian Karelia, and in Oslo fjord and Sognefjord in Norway. Four geographically vicariant sister clades were observed on graylings: Clade III in the Baltic Sea Basin; Clade IV in Karelian rivers draining to the White Sea; Clade V in Norwegian river draining to Swedish lake V?nern; and Clade VI in rivers draining to Oslo fjord. The pattern fitted perfectly with the postglacial history of grayling distribution. Widely sampled clades from salmon and Baltic grayling had basal haplotypes in populations, which were isolated early during the postglacial recolonisation. The divergence between the six clades was clear and linked with their hosts, but not wide enough to support a species status for them. Parasites from the Slovakian type population of G. thymalli were not available, so this result does not mean that G. salaris and G. thymalli are synonyms. It is suggested that the plesiomorphic host of the parasite cluster was grayling, and the switch to salmon occurred at least once when the continental ice isolated Baltic salmon in an eastern freshwater refugium, 130,000 years ago. At the same time, parasites on grayling were split geographically and isolated into several allopatric refugia. The divergence among the parasite clades allowed a tentative calibration of the evolutionary rate, leading to an estimate of the divergence of 13.7-20.3% per million years for COI coding mtDNA. The results supported the hypothesis that parallel to the allopatric mode, host switch and instant isolation by host specificity can be operated as a speciation mechanism.     

Identification of genetic markers associated with Gyrodactylus salaris resistance in Atlantic salmon Salmo salar

Gyrodactylus salaris Malmberg, 1957 is a freshwater monogenean ectoparasite of salmonids, first recorded in Norway in 1975 and responsible for extensive epizootics in wild Atlantic salmon Salmo salar L. The susceptibility of different populations of Atlantic salmon to G. salaris infection differs markedly, with fish from the Baltic being characterised as relatively resistant whereas those from Norway or Scotland are known to be (extremely) susceptible. Resistance to Gyrodactylus infection in salmonids has been found to be heritable and a polygenic mechanism of control has been hypothesised. The current study utilises a 'Quantitative trait loci' (QTL) screening approach in order to identify molecular markers linked to QTL influencing G. salaris resistance in B1 backcrosses of Baltic and Scottish salmon. Infection patterns in these fish exhibited 3 distinct types; susceptible (exponential parasite growth), responding (parasite load builds before dropping) and resistant (parasite load never increases). B1 backcross fish were screened at 39 microsatellite markers and single marker-trait associations were examined using general linear modelling. We identified 10 genomic regions associated with heterogeneity in both innate and acquired resistance, explaining up to 27.3% of the total variation in parasite loads. We found that both innate and acquired parasite resistance in Atlantic salmon are under polygenic control, and that salmon would be well suited to a selection programme designed to quickly increase resistance to G. salaris in wild or farmed stocks.     

An agent-based modelling approach to estimate error in gyrodactylid population growth

Comparative studies of gyrodactylid monogeneans on different host species or strains rely upon the observation of growth on individual fish maintained within a common environment, summarised using maximum likelihood statistical approaches. Here we describe an agent-based model of gyrodactylid population growth, which we use to evaluate errors due to stochastic reproductive variation in such experimental studies. Parameters for the model use available fecundity and mortality data derived from previously published life tables of Gyrodactylus salaris, and use a new data set of fecundity and mortality statistics for this species on the Neva stock of Atlantic salmon, Salmo salar. Mortality data were analysed using a mark-recapture analysis software package, allowing maximum-likelihood estimation of daily survivorship and mortality. We consistently found that a constant age-specific mortality schedule was most appropriate for G. salaris in experimental datasets, with a daily survivorship of 0.84 at 13°C. This, however, gave unrealistically low population growth rates when used as parameters in the model, and a schedule of constantly increasing mortality was chosen as the best compromise for the model. The model also predicted a realistic age structure for the simulated populations, with 0.32 of the population not yet having given birth for the first time (pre-first birth). The model demonstrated that the population growth rate can be a useful parameter for comparing gyrodactylid populations when these are larger than 20-30 individuals, but that stochastic error rendered the parameter unusable in smaller populations. It also showed that the declining parasite population growth rate typically observed during the course of G. salaris infections cannot be explained through stochastic error and must therefore have a biological basis. Finally, the study showed that most gyrodactylid-host studies of this type are too small to detect subtle differences in local adaptation of gyrodactylid monogeneans between fish stocks.     

Characterisation of a low pathogenic form of Gyrodactylus salaris from rainbow trout

Gyrodactylus salaris was isolated from rainbow trout in a Danish freshwater trout farm, and a laboratory population of this particular parasite form was established on rainbow trout. Challenge infections were performed using different salmonid strains and species, including East Atlantic salmon Salmo salar (from the Danish River Skjern?), Baltic salmon S. salar (from the Swedish River Ume Alv) and rainbow trout Oncorhynchus mykiss (from the Danish rainbow trout farm Fousing). These were compared to infection studies on the Norwegian Laerdalselva parasite form kept under exactly the same conditions in the laboratory. The Danish G. salaris form had low virulence towards both Atlantic and Baltic salmon, whereas rainbow trout proved susceptible to the parasite. The Danish G. salaris form was able to maintain a very low infection on East Atlantic salmon, but not on the Baltic salmon, which eliminated the infection within 2 wk. Rainbow trout developed infection intensities ranging up to several hundred parasites per host. The host colonization patterns of the parasite differed clearly from those of previous studies on microhabitats of the Norwegian form of G. salaris. A comparative study on morphological characters (opisthaptoral hard parts) from the Danish parasite form and Norwegian G. salaris showed no significant differences. Selected genes comprising internal transcribed spacers 1 and 2 (ITS), ribosomal RNA intergenic spacer (IGS) and cytochrome c oxidase subunit I (COI) regions were cloned and sequenced. Five sequenced ITS clones from 5 individuals of the Danish strain consistently revealed a single base substitution compared to ITS sequences from all other known species and strains of Gyrodactylus. Mitochondrial COI gene sequences demonstrated that the Danish G. salaris form is closely similar to the Laerdalselva parasite form found in Norway. The IGS sequences were highly variable, but very similar to those obtained from German isolates of G. salaris.     

Nestedness in assemblages of gyrodactylids (Monogenea: Gyrodactylidea) parasitising two species of cyprinid--with reference to generalists and specialists

The structure of gyrodactylid assemblages in individual fishes of two species of cyprinid was determined. A total of 100 specimens of minnow, Phoxinus phoxinus, and 137 specimens of roach, Rutilus rutilus, were investigated for presence of gyrodactylids. Host specificity, specialists vs. generalists, was noted in each host fish. A nested pattern was recorded in parasite assemblages of minnow, the host with a dominant number of specialist gyrodactylids. A non-nested pattern was observed in parasite assemblages of roach, the host with a dominant number of generalist gyrodactylids. The host specificity appears to be a meaningful factor that determines the pattern of gyrodactylid assemblages of both fish hosts.     

Analysis of ribosomal RNA intergenic spacer (IGS) sequences in species and populations of Gyrodactylus (Platyhelminthes: Monogenea) from salmonid fish in Northern Europe

The intergenic spacer (IGS) region of ribosomal RNA genes was amplified and sequenced from a variety of Gyrodactylus specimens collected from wild and farmed Atlantic salmon Salmo salar, rainbow trout Oncorhynchus mykiss, and grayling Thymallus thymallus, from various locations in Northern Europe. Phylogenetic analysis of the sequences confirmed the distinction between G. salaris Malmberg, 1957 and G. thymalli Zitnan, 1960, supporting their validity as separate species. G. salaris adapted to rainbow trout are also distinct from the parasites found on Atlantic salmon, supporting the existence of a rainbow-trout form that was initially identified on the basis of morphological differences. Analysis of the IGS did not provide good resolution of different populations of G. salaris sensu stricto, but was consistent with epidemiological evidence which indicates that introduction of the parasite to Norway was recent and limited. The IGS may be helpful in distinguishing forms of G. salaris that are pathogenic to Atlantic salmon from those that are not.     

Differences in susceptibility of anadromous and resident stocks of Arctic charr to infections of Gyrodactylus salaris, under experimental conditions

The ability of Gyrodactylus salaris , an important pathogen of the Atlantic salmon Salmo salar , in Norway, to infect anadromous and resident stocks of the Arctic charr, Salvelinus alpinus , has been examined in the laboratory. Resident charr (Korssjoen stock) exposed to heavily infected salmon, were considered innately resistant as they lost their infections within 21 days when individually isolated. Isolated anadromous harr (Hammerfest stock) remained infected for up to 150 days, although most infections disappeared within 30–50 days. In many cases the parasite population grew initially, but growth was limited after 20–30 days and infections subsequently disappeared. At the same time, shoals of 50 anadromous charr, swimming in the tanks containing the individually isolated fish in floating cages, remained infected for up to 280 days. Charr isolated from these shoals after 115 days and subsequently monitored individually lost their infections within 30 days, although the parasite persisted within the shoals for a further 75–135 days. This suggests that G. salaris , persisted on shoaling charr despite an immune response which led to the elimination of parasites from isolated hosts. The Hammerfest stock of anadromous charr supports G. salaris , in the laboratory, and the extended period of survival on this host suggests that charr may be important in the epidemiology of G. salaris , in northern Norway.     

Susceptibility of Baltic and East Atlantic salmon Salmo salar stocks to Gyrodactylus salaris (Monogenea)

The susceptibility of a Baltic salmon stock Salmo salar (Indals?lv, central Sweden) to Norwegian Gyrodactylus salaris (Figga strain, central Norway) was experimentally tested and compared with previously obtained results on East Atlantic salmon (Lierelva, SE Norway). Contrary to expectation, the Baltic salmon, which had no prior exposure to this parasite strain, appeared almost as susceptible as the Norwegian salmon parr that naturally experience G. salaris-induced mortality. Individually isolated salmon of both stocks sustained G. salaris infections with little evidence of innate resistance. A few individuals of the Indals?lv stock controlled their infection from the beginning, but overall there was considerable heterogeneity in the course of infection in both stocks. On individual hosts, G. salaris growth rates declined steadily throughout the infection, a trend which was particularly marked amongst the Lierelva stock. On shoaling Lierelva fish, there was some evidence of reduced parasite population growth towards the end of the infection; this was not apparent in Indals?lv fishes. These results reflect a growing awareness that not all Baltic salmon may be resistant to Norwegian G. salaris, and that Norwegian and Baltic G. salaris strains may differ in virulence. Consequently, management decisions concerning this parasite-host system should be based upon the actual, and tested, susceptibility of stocks under consideration and not upon identification of stocks as either Atlantic or Baltic.     

Gyro-scope: an individual-based computer model to forecast gyrodactylid infections on fish hosts

Individual-based computer models (IBM) feature prominently in current theoretical ecology but have only been applied in a small number of parasitological studies. Here we designed an IBM to simulate the infection dynamics of gyrodactylid parasites and immune defence of na?ve hosts (i.e. fish previously not exposed to these parasites). We compared the results of the model with empirical data from guppies (Poecilia reticulata) infected with Gyrodactylus parasites. The laboratory experiments on guppies showed that larger fish acquired a heavier parasite load at the peak of the infection. The survival probability declined with increased body size and no fish survived a parasite load of 80 or more worms in this experiment (i.e. lethal load). The model was a good predictor of the Gyrodactylus infection dynamics of guppies and the model output was congruent with previously published data on Gyrodactylus salaris infections of salmon (Salmo salar). Computer simulations indicated that the infections persisted longer on larger hosts and that the parasite load increased exponentially with the body size of the host. Simulations furthermore predicted that the parasite load of fish with a standard length in excess of 17mm (i.e. the size of adult guppies) reached a lethal load. This suggests that in the conditions of the experiment, the immune defence of na?ve guppies can offer moderate protection against gyrodactylid infections to juveniles, but not to na?ve adult guppies. The model is a useful tool to forecast the development of gyrodactylid infections on single hosts and make predictions about optimal life history strategies of parasites.     

Mode of transmission, host switching, and escape from the Red Queen by viviparous gyrodactylids (Monogenoidea)

Compared to other monogenoidean groups, viviparous gyrodactylids exhibit extraordinary species diversity and broad host range. It has been suggested that this evolutionary success is associated with a suite of morphological and life-history traits that include, in part, continuous transmission (i.e., ability to infect new hosts throughout the gyrodactylid life cycle). Experiments were conducted to explore the putative adaptive advantage of continuous transmission within viviparous gyrodactylids during colonization of new host resources. Differences in infrapopulation growth, such as abundance, prevalence, and duration of the infection, of Gyrodactylus anisopharynx on 3 species of fish--Corydoras paleatus and Corydoras ehrhardti (both natural hosts) as well as Corydoras schwartzi (a host not known to harbor G. anisopharynx)--held under isolated and grouped conditions were determined. Results showed that infrapopulations of G. anisopharynx on C. paleatus and C. schwartzi had higher growth when the parasite had the opportunity for host transfer (grouped hosts). Infrapopulations of G. anisopharynx on isolated and grouped C. ehrhardti showed an opposite trend, although differences in mean duration and maximum abundance were not statistically different. Results obtained from experiments with C. paleatus and C. schwartzi support the hypothesis that continuous transmission in viviparous gyrodactylids enhances colonization success, probably by allowing initial avoidance of Red Queen dynamics. The absence of statistical differences between infrapopulations on isolated and grouped C. ehrhardti suggests that parasite dynamics may be influenced by factors other than continuous transmission in this host.     

The ultrastructure of hypersymbionts on the monogenean Gyrodactylus salaris infecting Atlantic salmon Salmo salar

There is increasing pressure to develop alternative control strategies against the pathogen Gyrodactylus salaris, which has devastated wild Atlantic salmon Salmo salar in Norway. Hyperparasitism is one option for biological control and electron microscopy has revealed two ectosymbionts associated with G. salaris: unidentified rod-shaped bacteria, and the protist, Ichthyobodo necator. No endosymbionts were detected. The flagellate I. necator occurred only occasionally on fish suffering costiosis, whereas bacterial infections on the tegument of G. salaris were observed throughout the year, but at variable densities. Bacteria were seldom observed attached to fish epidermis, even when individuals of G. salaris on the same host were heavily infected. Wounds on salmon epidermis caused by the feeding activity of bacteria-infected G. salaris did not appear to be infected with bacteria. On heavily infected gyrodactylids, bacteria were most abundant anteriorly on the cephalic lobes, including the sensory structures, but no damaged tissue was detected by transmission electron microscopy in the region of bacterial adherence. Furthermore, transmission and survival of infected G. salaris on wild salmon did not appear to be influenced by the bacterial infection. The lack of structural damage and impact on G. salaris biology indicates that these bacteria are not a potential agent for control of gyrodactylosis. However, this may not be the case for all gyrodactylid-bacterial interactions and a review of bacterial infections of platyhelminths is presented.     

Dexamethasone treatment affects skin mucous cell density in Gyrodactylus derjavini infected Salmo salar

Atlantic salmon, Salmo salar, is normally rather refractive to infection with the ectoparasitic monogenean Gyrodactylus derjavini but dexamethasone treatment of the host increases the susceptibility. The causative mechanisms were elucidated in this work. Groups of Atlantic salmon were treated by intra-peritoneal dexamethasone injections and subsequently infected with G. derjavini. It was shown that both the infection level and the mucous cell density of caudal and pelvic fins were affected by the treatment. Significantly higher mucous cell densities were found on infected and treated fish whereas non-infected and treated fish showed no significant elevation of cell density. This suggests that mucous cell discharge elicited by infection is inhibited by the drug. The association with elevated parasite counts in these fish can be explained either by decreased anti-parasitic mucus action or by parasite predilection for intact mucous cells.     

Selection by parasites in spate conditions in wild Trinidadian guppies (Poecilia reticulata)

Gyrodactylids are ubiquitous fish parasites and yet, with the notable exception of Gyrodactylus salaris, few studies have reported the effect of these parasites on host survival in natural populations. Here, we assess the impact of the parasite load of gyrodactylids (G. turnbulli and G. bullatarudis) on the survival and migration of guppies (Poecilia reticulata) in their natural habitat of the Aripo River in Trinidad. The recapture rate of males declined by 19% with every additional parasite, a remarkably high figure given that the parasite load in this study ranged from zero to 20 worms. In addition, with an increased number of parasites, males were more prone to be recovered downstream. In contrast, no effect of parasitism was observed in females. The mean parasite load sharply declined after a series of flushing events during heavy seasonal downpours. The parasite load varied significantly between fish depending on their location in the river, and the size of the fish explained variation in parasite load between individuals. The present study indicates that tropical gyrodactylid parasites can play an important role in the ecology of natural fish populations, causing intense bouts of natural selection in guppies during heavy rains in the wet season.     

Comparative susceptibility of two races of Salmo salar (Baltic Lule river and Atlantic Conon river strains) to infection with Gyrodactylus salaris

The susceptibility of various races of salmonids towards infections with the skin parasitic monogenean Gyrodactylus salaris Malmberg, 1957, differs markedly. Norwegian and Scottish salmon strains are known as extremely susceptible to infection, whereas Baltic salmon races such as the Neva strain (Russian origin) and the Indals river (Swedish origin) salmon have been characterized as relatively resistant. However, the status of the many other Baltic strains has remained unknown. The present study reports on the susceptibility of the Baltic salmon from the Swedish river Lule. It was shown that this strain is susceptible to infection but to a lesser extent than the Scottish salmon. Further studies showed that injection of immuno-suppressants (dexamethasone) greatly increased population growth of G. salaris on Scottish salmon but not on the Baltic salmon. Mucous cell density on fins differed between strains, and a general trend to decreased cell density on infected fish 8 wk post-infection, compared to uninfected fish, was observed. The largest decrease in mucous cell density following infection was seen in the most resistant fish. After administration of immuno-suppressants, this decrease in mucous cell density was inhibited in the Scottish salmon but not in the Baltic salmon. Thus, there seems to be a relationship between the fishes' ability to discard mucous cells and the ability to resist infections with Gyrodactylus salaris. Although the Lule salmon seems more susceptible to infection compared to previous reports on the Neva salmon, the results support the notion that Baltic salmon strains are generally more resistant than East Atlantic salmon.     

Molecular faunistics of accidental infections of <Emphasis Type="Italic">Gyrodactylus</Emphasis> Nordmann, 1832 (Monogenea) parasitic on salmon <Emphasis Type="Italic">Salmo salar</Emphasis> L. and brown trout <Emphasis Type="Italic">Salmo trutta</Emphasis> L. in NW Russia

Salmon Salmo salar L. and brown trout S. trutta L. juveniles were examined for the presence of accidental monogenean ectoparasitic species of Gyrodactylus Nordmann, 1832 in the Baltic and White Sea basins of Russian Karelia in order to estimate the frequency of host-switching attempts on an ecological timescale. To collect phylogeographical information and for exact species identification, the parasites were characterised by nuclear internal transcribed spacer sequences of rDNA (ITS) and, for some species, also by their mitochondrial DNA (CO1 gene) sequences. Four accidental Gyrodactylus species were observed on salmon and brown trout. A few specimens of G. aphyae Malmberg, 1957, the normal host of which is the Eurasian minnow Phoxinus phoxinus (L.), were observed on lake salmon from the Rivers Kurzhma (Lake Kuito, White Sea basin) and Vidlitsa (Lake Ladoga, Baltic basin). G. lucii Kulakovskaya, 1952, a parasite of the northern pike Esox lucius L., was observed on salmon in the Kurzhma. In the River Vidlitsa, two specimens of G. papernai Ergens & Bychowsky, 1967, normally on stone loach Barbatula barbatula (L.), were found on salmon. On anadromous White Sea salmon in the River Pulonga in Chupa Bay, a few salmon parr carried small colonies of G. arcuatus Bychowsky, 1933, which were shown to have originated from the local three-spined stickleback Gasterosteus aculeatus L. consumed as prey. No specimens of Gyrodactylus salaris Malmberg, 1957 were observed, although the Pulonga is the nearest salmon spawning river to the River Keret', which is heavily infected with introduced G. salaris. In the River Satulinoja, Lake Ladoga, three specimens of G. lotae Gusev, 1953, from burbot Lota lota (L.), were collected from a single brown trout S. trutta. All nonspecific gyrodactylid infections on salmonids were judged to be temporary, because only a few specimens were observed on each of the small number of infected fishes. The prevalence of endemic G. salaris was also low, only 1% (Nfish = 296) in Lake Onega and 0.7% (Nfish = 255) in Lake Ladoga, while brown trout specific Gyrodactylus species were not observed on any of the 429 trout examined from the Ladoga basin. The host-specific and unspecific burden of Gyrodactylus spp. on these 'glacial relict' populations of salmon and brown trout was very low, suggesting a generalised resistance against the co-evolved freshwater parasite community, or some kind of 'vaccination' effect. These hypotheses deserve further testing.