The dynamics of infection of Diphyllobothrium dendriticum (Nitzsch) and D. ditremum (Creplin) in the char Salvelinus alpinus (L.) in Sweden

A total of 654 char, Salvelinus alpinus (L.), from Lake Bjellojaure, Sweden, were examined for plerocercoids of Diphyllobothrium dendriticum (Nitzsch, 1824) and D. ditremum (Creplin, 1825) from September 1969 to December 1974. Plankton samples showed that the dominating copepod species in the lake were Eudiaptomus graciloides (Lilljeborg), Cyclops scutifer Sars and Heterocope appendiculata Sars. E. graciloides passed the winter as adults and C. scutifer as copepodids while H. appendiculata was restricted to a short period in the summer. Copepods occurred in the stomachs in September-October and to a certain extent in April-May, but only cyclopids and Heterocope sp. were found. C. scutifer is proposed to be the main first intermediate host of both D. dendriticum and D. ditremum . The potential final hosts were thecommon gull, Larus canus L., for D. dendriticum and the black-throated diver, Gavia arctica (L.), for D. ditremum . One common gull was experimentally successfully infected with plerocercoids of D. dendriticum . The parasites showed no seasonal cycle in incidence of infection.     

Egg size and form as taxonomic criteria in Diphyllobothrium (Cestoda, Pseudophyllidea)

The size and form (length, width, and length: width ratio) of eggs of Diphyllobothrium dendriticum, D. ditremum and D. latum vary considerably among individual worms within each species. The size of eggs varies with host species and a decrease in egg size with increasing intensity of infestation is indicated. The egg size of D. latum increases during the first 10-12 days of egg production. For single worm infections in golden hamsters the mean egg length and width of D. ditremum are significantly smaller than the corresponding means of D. dendriticum and D. latum, while D. latum has significantly wider eggs than D. dendriticum. As taxonomic characteristics, egg size and form may contribute to species delimitation at the population level. For identification at the individual level the best possible accuracy is about 80%. This accuracy is considerably reduced when variation in host species and intensities of infestations are introduced. Scanning electron microscope studies did not reveal any differences among eggs of the three species.     

Aggregation of helminths: the role of feeding behavior of fish hosts

Individual Arctic charr (Salvelinus alpinus) from Fjellfr?svatn, northern Norway, could be categorized by their stomach contents as zooplanktivores or benthivores. Feeding specialization among these fish was evident from negative correlations between helminths transmitted by pelagic copepods (Diphyllobothrium dendriticum and D. ditremum) and those transmitted by the benthic amphipod Gammarus lacustris (Cystidicola farionis and Cyathocephalus truncatus). Occurrences of parasite species acquired from the same types of invertebrate were positively correlated in the fish. Strong relationships among habitat use, diet, and helminth infections among the Arctic charr indicated persistent foraging patterns involving long-term habitat use and feeding specialization. The distribution of all parasite species was highly aggregated in the fish samples, measured by the exponent k of the fitted negative binomial distributions (range: 0.5-7.5) and the variance-to-mean ratios (s2/mean, range: 5-85). Charr specializing on either copepods or Gammarus predominantly contributed to high-intensity class intervals within the overall frequency distributions of the corresponding parasite species. Such fish had low infection intensities of helminths transmitted by other prey organisms. The detailed analyses of the parasite frequency distributions for fish with different habitat or feeding preferences evidently show how heterogeneity in trophic behavior contributes strongly to the commonly observed aggregation of helminths among hosts under natural conditions.     

Ecological parasitology of polymorphic Arctic charr, Salvelinus alpinus (L.), in Thingvallavatn, Iceland

The helminth endoparasite fauna in four Arctic charr morphs, Salvelinus alpinus (L.), small benthivorous (SB), large benthivorous (LB), planktivorous (PL) and piscivorous (PI) charr, from Thingvallavatn, Iceland consisted of: Crepidostomum farionis (Trematoda: Allocreadiidae); Diplosttomum sp. (Trematoda: Diplostomatidae); Eubothrium salvelini; Diphyllobothrium dendriticum; D. ditremum (Cestoda: Pseudophyllidae); Proteocephalus longicollis (Cestoda: Proteocepha-lidae): and Philonema oncorhynchi (Nematoda: Filariidae). The morphs exhibited distinctive patterns in prevalences and parasite burdens (mean intensity and mean relative density of parasites). SB charr had high prevalence and parasite burden of the eye fluke Diplostomum sp. and none to very light infections of the other parasite species. LB charr had relatively high prevalence and parasite burden of the intestinal fluke C. farionis , whereas infections of the remaining parasite species were light to moderate. PL and PI charr had high prevalences and worm burdens of Diphyllobothrium spp. and P. longicollis . PL charr differed from PI charr in higher worm burden off P. longicollis and lighter burden of £. salvelini . Prevalences of P. oncorhynchi were high in PL and PI charr. Association of parasite intensities and age and length offish were investigated. The different infection patterns among the morphs agree well with their partitioning in food and habitat utilization, and confirm that there is a high degree of ecological segregation between the morphs. The results demonstrate the importance of ecological factors influencing transmission efficiency of parasites to the fish host.     

Parasites as indicators of individual feeding specialization in Arctic charr during winter in northern Norway

The relationship between infection with the food-transmitted parasites Diphyllobothrium dendriticum, D. ditremum (Cestoda) and Cystidicola farionis (Nematoda) and prey selection was studied in individual Arctic charr from Lake Takvatn, northern Norway. There was no correlation between parasites transmitted throughout prey organisms from benthic habitats (amphipods) and pelagic habitats (copepods). A strong relationship between infection with a parasite species and the corresponding intermediate host from the stomach content of individual charr, indicated an individual feeding specialization. Independent of size, charr specialized on the intermediate hosts of all three parasite species. Some charr maintained this specialization on specific prey items throughout the winter period. These parasite species are considered to be useful indicators of past prey selection.     

The ecological interaction between arctic charr, Salvelinus alpinus (L.), and the plerocercoid stage of Diphyllobothrium ditremum

Six hundred and two arctic charr, Salvelinus alpinus , from a small lake (30 ha) were examined for infection with plerocercoids of Diphyllobothrium ditremum the only species of this genus present; 466 (77.4%) harboured the parasite in an overdispersed infection. The prevalence of infection increased with age and reached 100% at 7+ years for males and 8+ years for females. The rate of infection appeared to be related to the feeding biology of the charr which did not change within the age span studied. The rate of increase in density of infection (plerocercoids per fish) decreased with age as did the degree of overdispersion. It is concluded that the infection probably induces mortality in the fish population, and that such mortality is dependent on parasite density.     

Geographical distribution and host-parasite relationships of Plistophora ovariae (Microsporida, Nosematidae) in Notemigonus crysoleucas

Plistophora ovariae undergoes schizogony and sporogony in developing ova of the golden shiner. Destruction of the ova greatly reduces fecundity and causes spawning failures. The incidence and intensity of infection is described in 49 commercial fish farms from 12 states. The parasite was found in fish from 45 of 49 sources and in a farm pond and creek from Oklahoma. Overall incidence of infection was 48% of 2759 fish. No significant difference was found in the incidence of disease from fish propagated by either intensive or extensive culture methods. There was a significant difference in incidence of infection with host age. Incidence increased from 30% in age class 0 to 75% in age class 4. Intensity of infection decreased with age and varied with season; it was greatest in May and June. Thus, the maximum number of spores and infected ova occurred during the spawning season of the host. Infected fish were generally larger and heavier than uninfected fish. Reduced egg production (partial parasitic castration) allowed nutrients and energy to be used for faster growth.     

Antigenic affinity of species of the genus Diphyllobothrium (Cestoidea, Diphyllobothriidae) distributed in the territory of Karelia

Experiments on ringprecipitation reactions with homologous and heterologous antigenes of four species of diphyllobothriids have shown that D. dendriticum and D. ditremum are in a more close antigenic affinity. In relation to the above two species D. vogeli is approximately at the same level. D. latum displayed no antigenic affinity with the above species that, apparently, is connected with its parasitism only in mammals. Schemes of immunization of rabbits for obtaining antisera are given.     

Ultrastructural studies on Diphyllobothrium ditremum and D. dendriticum (Cestoda, Pseudophyllidea), with emphasis on the scolex tegument and the tegument in the area around the genital atrium.

The scolices of D. ditremum and D. dendriticum are similar in regard to general morphology and ultrastructure of the tegument. At the scolex apex and along the upper bothrial edges sensory endings are numerous. The inner bothrium surface of D. ditremum appears to be plain or smooth, the distal cytoplasm is only about 0.6 mum thick and covered with long and slender microtriches. When D. ditremum is attached to the host gut an adhesive layer of secreted substance exists between the inner bothrium surface and the host intestinal villi. In D. dendriticum, however, no secreted layer exists between the inner bothrium surface and the host intestinal villi and the inner bothrium surface of this species appears to be lobed or lappet formed. The lobes are protrusions of the distal cytoplasm and are covered with long and slender microtriches. Large conglomerations of secretion globules appearing to be enclosed by a double membrane, are observed in the scolices of D. dendriticum and D. ditremum, although more frequent in the latter. Differences existing between the distal cytoplasm of the scolex and the gravid proglottid are described. The papillae around the genital atrium in mature and gravid segments were studied. Each papilla appears as a thickening of the distal cytoplasm, which in this region is 10-12 mum thick, when studied with transmission electron microscope. From the area between the papillae sensory endings appearing in sections to be either single, double or triple are described.     

The distribution and biology of the cestode Eubothrium parvum in capelin, Mallotus villosus, (Pallas) in the Barents Sea, and its use as a biological tag

Samples of Eubothrium parvum were obtained from capelin Mallotus villosus at 55 stations throughout the Barents Sea and from Balsfjord, North Norway. The parasite is distributed widely throughout the Barents Sea, but both incidence and intensity of infection are higher in the regions off Murmansk and the Kola peninsula, and Spitsbergen. E. parvum exhibits a seasonal peak in maturation and probably also in acquisition of new infections. The incidence of infection is greatest in 1 + fish, whereas the intensity is more independent of host age. It is suggested that the parasite requires only a single intermediate host, a plank-tonic copepod, and its distribution in relation to age of host is a reflection of the dietary preference shown by young capelin for copepods. The frequency distribution of E. parvum in capelin was over-dispersed in Balsfjord, where infection levels of between 1 and 28 parasites per fish were encountered in all samples, but under-dispersed in the Barents Sea, where infections of more than four parasites per fish were never found and even infections with three and four parasites were very local. It is suggested that the underdispersion is due to a very low probability of infection in the open waters of the sea. Although the presence of E. parvum cannot be used as a biological tag for capelin, its abundance and frequency distribution can. The difference in frequency distribution and the failure to find any heavily infected fish in the Barents Sea confirm the suggestion that the capelin of Balsfjord form a local isolated population, which does not migrate into the Barents Sea. The differences in infection levels within the Barents Sea suggest the further possibility that there are at least two stocks of capelin there, but this requires further investigation and confirmation.     

The population biology of two species of eyefluke, Diplostomum gasterostei and Tylodelphys clavata, in perch

Incidence, intensity and frequency distributions of infections in perch, Perca fluviatilis , with the eyeflukes Tylodelphys clavata and Diplostomum gasterostei were studied in a lake at monthly intervals over a period of two years. T. clavata had appeared in the lake immediately prior to the study, and its levels of infection continued to rise throughout the period of investigation whereas those of D. gasterostei remained relatively steady. The main period of infection of fish by both species of parasites was late summer, but a second and minor infection took place in spring. In the first year after its introduction T. clavata exhibited a seasonal cycle in incidence and intensity of infection and disappeared from fish in summer, but in the second year this cycle was masked by increased variation in development times and lack of synchrony throughout the parasite population. It is concluded that T. clavata has a life span of one year or less and that the parasites die within the fish, mainly in summer. D. gasterostei in contrast has a life span of at least one year, and disappearance of the parasite from the population is probably due to death of some heavily infected fish, although it was not possible to determine if this was due directly to the presence of the parasite.     

The population biology of the acanthocephalan Pomphorhynchus laevis (Müller) in the River Avon

Regular samples of Gammarus pulex and dace Leuciscus leuciscus and occasional grayling Thymallus thymallus and chub L.cephalus were examined from the River Avon, Hampshire, for the presence of the acanthocephalan Pomphorhynchus leavis . The parasite only occurred in medium sized Gammarus due to lower probability of contact with small gammarids and stunted growth and selective mortality amongst older infected ones. No cycles in incidence or development of the parasite in G.pulex were observed. The parasite infected gammarids and grew in all months, and cystacanths were available throughout the year. Despite seasonal feeding activity and dietary preferences, fish fed on Gammarus and acquired infections in all months. Dispersion of P.laevis within the fish population was related to host feeding behaviour. No evidence of seasonal cycles in incidence or intensity of infection in fish was found, and observed monthly changes in the parasite population were related to changes in size structure of the host sample. In dace and grayling P.laevis grew little and matured only in summer, but in chub it grew and produced acanthors all year. The parasite population in fish appeared to be in a state of dynamic equilibrium and gain and loss of parasites took place throughout the year with the level of infection at any moment being determined primarily by the feeding behaviour of the host. This relationship between host diet, water temperature and parasite population size is discussed, and P.laevis in the R. Avon compared with other localities and other parasites.     

Distribution of the cestode Caryophyllaeides fennica (Cestoda: Lytocestidae) in the population of cyprinid fish in the Rybinsk water reservoir

The distribution of the cestode Caryophyllaeides fennica in populations of the ide, roach, white bream and bream from the Rybinsk reservoir has been analyzed in regard to fish age. The relative abundance of C. fennica in populations of each host species was calculated as product of the helminthes abundance by the fish number in age groups. The highest prevalence and abundance of cestodes was found in the ide. All age classes of this species were infected, with the maximum in fishes of the age 3+ ... 5+. The roach is infected with C. fennica till only 10-year age; brean is infected till the age 4+. In the white bream C. fennica is an occasional parasite. The cestode number among hosts was as follows: bream--68%; roach--26%, ide--5%, white bream--1%. Different approaches to the estimation of the parasite abundance distribution among several host species (in terms of mean prevalence and intensity of the relative abundance of parasites) are discussed.     

Infection of estuarine fish populations by Cryptocotyle lingua (Creplin)

Fish populations in the Medway Estuary become infected with Cryptocotyle lingua (Creplin) shortly after their inshore migrations. The present study examines infection levels between 1973 and 1975 in four fish species; whiting, Merlangius merlangus (L.), pouting, Trisopterus luscus (L.), plaice, Pleuronectes platessa L. and flounder, Platichthys flesus (L.). Frequency of infection was highest in flounder, followed by pouting, plaice and whiting. Unusually, parasite density was lowest in plaice even though gastropod molluscs (the parasite's first intermediate host) were most common in their diet. Seasonal levels of infection are discussed and prove useful indicators to the migratory movements of individual fish populations. In whiting and pouting, incidence of infection increased with length of host while no relationship between infection and age of host was observed in either flatfish. Metacercariae were not distributed randomly over the body surface; in gadoids they were concentrated above the lateral line while in pleuronectids they were most intense in the fins and gills.     

Field and experimental studies on Dicrocoelium dendriticum and dicrocoeliasis in northern Spain

The transmission, control and the relationship between Dicrocoelium dendriticum and its definitive (sheep and cattle) and intermediate (molluscs and ants) hosts under natural and experimental conditions are described. Eleven species of molluscs and four of ants were found infected with larval D. dendriticum in León province, north-west Spain. Infected ants were observed between April and November and in tetania at 7.5-26.9 degrees C. The highest shedding of eggs by sheep and cattle was detected in winter. Two treatments applied in November and January were the most effective. In experimentally infected molluscs, the parasite was not visible under the stereomicroscope, at least until 50 days post-infection (p.i.). The prepatent period in experimentally infected lambs was 49-79 days p.i. The number of eggs per gram increased with the days p.i. and the parasite burden. The aspartate aminotransferase, alanine aminotransferase, gamma-glutamyl transpeptidase, leukocyte and neutrophil values of infected lambs increased, but those of lymphocytes decreased. Using the enzyme-linked immunosorbert assay technique, the IgG antibody response to excretory-secretory and somatic antigens of D. dendriticum was positive from day 30 p.i., although the maximum antibody levels were observed on day 60 p.i. The number of worms per lamb ranged between 30 and 2063. Cholangitis and cholangiectasia of the septal bile and hepatic ducts were observed. The best enzymatic systems for adult and larval D. dendriticum characterization were lactate dehydrogenase, glucose phosphate isomerase and phosphoglucomutase. Genetic variability of adult D. dendriticum was high using the random amplified polymorphic DNA technique.     

Seasonal dynamics in abundance, development and pattern of infection of Bunodera luciopercae (Müller) in perch, Perca fluviatilis L. from an oligotrophic lake in Norway

Samples of perch, Perca fluviatilis L., were examined monthly for the presence of the intestinal trematode Bunodera luciopercae (Müller). Marked seasonal cycles in incidence and intensity of infection were found, with a minimum level in July, when all the fish examined were free of the fluke, and a maximum level in February. Five developmental stages of the fluke were recognized and the seasonal changes in the percentage distribution of these stages were studied. Infective larvae were found in the period August–March, with a peak in January. Gravid worms were mainly found in the period May-June. The frequency distribution of parasite counts were over-dispersed and followed the negative binomial distribution within fish samples from different periods of the year.     

Populations of Lamproglena pulchella von Nordmann 1832 (Copepoda: Eudactylinidae) in cyprinid fish in rivers with different pollution levels

A total of 512 chub ( Leuciscus cephalus L.) individuals sampled from four rivers with different pollution levels were examined for Lamproglena pulchella (von Nordmann 1832) prevalence and mean intensity. As a result of the L. pulchella mode of reproduction, the parasite populations showed a significant degree of aggregation. The results, excluding the effect of fish health conditions on the parasite infection, provide evidence of the influence of water quality on parasite prevalence. Parasite density increase in relation to fish length and age suggests a dependence of infection intensity to the exposure time on the fish.     

Helminth infections of wild boars (Sus scrofa) in the Bursa province of Turkey

The present study aimed to investigate the status of helminth infections in wild boars in the Bursa province of Turkey. For this purpose, during 2007-2008, 27 wild boars were necropsied and examined for helminths. Individual samples of tongue and diaphragm from 27 necropsied wild boars and an additional 22 tongue and diaphragm samples provided by hunters were examined by trichinoscopy and artificial digestion for Trichinella spp. larvae. Twenty animals (74%) were identified as being infected with at least one helminth species. Twelve species of helminths were detected, with the following prevalence rates: Metastrongylus apri (59%), Metastrongylus salmi (52%), Metastrongylus pudendotectus (52%), Dicrocoelium dendriticum (33%), Globocephalus urosubulatus (22%), Macracanthorhynchus hirudinaceus (19%), Gongylonema pulchrum (11%), Physocephalus sexalatus (7%), Trichuris suis (7%), Ascarops strongylina (4%), Hyostrongylus rubidus (4%) and Taenia hydatigena larvae (4%). Generally, lungworms were the predominant helminths. The highest mean abundance was observed for M. pudendotectus, and the lowest was determined for T. hydatigena larvae. Significant differences in the prevalence and intensity were found for D. dendriticum with respect to host age and sex, respectively. The mean intensity of M. pudendotectus was significantly influenced by the sex and age of the wild boars. This study is the first report describing the presence of M. salmi, M. pudendotectus, D. dendriticum, G. urosubulatus, M. hirudinaceus, P. sexalatus, A. strongylina and H. rubidus in wild boars in Turkey. All analysed muscle samples were negative for Trichinella spp. larvae.     

Response of Ig-positive cells to Enteromyxum scophthalmi (Myxozoa) experimental infection in turbot, Scophthalmus maximus (L.): A histopathological and immunohistochemical study

In recent years, a new parasite that causes severe losses has been detected in farmed turbot, Scophthalmus maximus (L.), in Northwestern Spain. Dead fish showed emaciation and cachexia caused by severe necrotizing enteritis that affected all portions of the digestive tract. The parasite was classified as a myxosporean and named Enteromyxum scophthalmi. This study was set up to gain insights into the immune response of fish against this parasitic infection. The kinetics of immunoglobulin positive (Ig+) cells in spleen, kidney and intestine from turbot experimentally infected with E. scophthalmi was studied. For evaluating both the progress of infection and the lesions induced by the parasite, we performed histopathological studies and for detecting Ig+ cells in situ we used an indirect immunohistochemical method. In fish exposed to the parasite, the number of Ig+ cells significantly increased in the intestine, the target organ of the parasite, whereas in spleen and kidney, haematopoietic organs where the parasite was not detected, the number of Ig+ cells decreased. Furthermore, the pattern of distribution of Ig+ cells changed in all three organs examined in recipient/infected fish compared with control fish (not exposed to the parasite). The results obtained in this study indicate that the infection by E. scophthalmi in turbot induced an immune response that involved changes in the number and distribution of Ig+ cells.     

Peculiarities of the parasitofauna of Atlantic salmon parr (Salmo salar L.), brown trout (Salmo truttae L.), and char (Salvelinus alpinus L.) in the Utsjoki River system (Northern Finland)

Data on the parasite diversity in Salmonidae fish parr from different parts of the Utsjoki River obtained during 1993-1995 and 2006-2007 are presented. Three fish species, Salmo salar L., S. truttae L., and Salvelinus alpinus L., were examined on the presence of helminthes. Twenty species of salmon parasites were found, the majority of which are the parasites with complicated life cycles. Infusorians C apriniana piscium, myxosporidia Chloromyxum januaricus and Myxobolus neurobius, metacercaria of the genera Diplostomum and Apatemon, and the nematode Raphidascaris acus larvae were the most numerous in salmon parasite fauna. Brown trout had the most number of specific parasite species, whereas char was infested by protozoan parasites only.