Study of fecundity of Bunodera luciopercae (Trematoda: Bunoderidae)

The seasonal dynamics of the maturation of trematode Bunodera luciopercae was investigated in 1992-1998. The actual fecundity of adult helminths from the ruff, perch, sander and volga pike-perch in the Rybinsk Reservoir was registered in the end of spring. The accumulation process of eggs reflected the host alteration and depended on the food ration, body length and sex of hosts.     

Enteric helminths of perch (Perca fluviatilis L.) and yellow perch (Perca flavescens Mitchill): stochastic or predictable assemblages?

Component communities of perch (Perca fluviatilis L) in Eurasia and the North American yellow perch (Perca flavescens Mitchill) were examined to determine the nature of their parasite communities. The scale of this investigation is continental and includes data collected across the distribution of each host species. Data were compiled from the literature and from 5 sample sites in North America. Four parasite species were found to occur frequently in the helminth component communities of P. flavescens. The cestodes Bothriocephalus cuspidatus and Proteocephalus pearsei, the digenean Crepidostomum cooperi, and the nematode Dichelyne cotylophora comprised a suite of species of which some or all occurred in most samples. Similarly, a group of 4 predictable parasite species was identified for P. fluviatilis in Eurasia, the digenean Bunodera luciopercae, the nematode Camallanus lacustris, the cestode Proteocephalus percae, and the acanthocephalan Acanthocephalus lucii. Specificity was not a requirement for predictability. Despite geographical isolation for millions of years, and different fish species interactions within and between continents, the predictability of these parasite assemblages indicates they are shaped by a biology, especially feeding patterns, common to both perch species. This is evidence that parasite assemblages comprised of nonhost-specific parasites in freshwater fishes are not merely stochastic assemblages but have key components that are predictable at this broad continental scale.     

Seasonal variations of the localization of the Bunodera luciopercae marites (Trematoda: Bunoderidae) in three species of perch fishes

Distribution of Bunodera luciopercae marites in the intestine of the fuff, sande and perch were studied in the Rybinsk Reservoir. For the fuff and perch fry, except three summer months, the more proportion of helminths was observed in the posterior region of the intestine. Because of a continuous passage of B. luciopercae in predatory fishes (sanders and adult perches), more proportion of trematodes was always observed in anterior region of the intestine. In all groups of hosts, 20-30% of total parasite number was registered in the middle intestine.     

Emergence of the mature trematode Bunodera luciopercae from infected fish under different light conditions

The elimination dynamics of the sexually mature trematode Bunodera luciopercae, a specific parasite of the perch Perca fluviatilis L. was studied using three photoperiods (L: D; L: L; D:D). Permanent illumination induced the helminth to leave fish. The diurnal rhythm of elimination is not pronounced. High values were recorded both in the morning and in the evening. A 40-h elimination rhythm revealed for B. luciopercae is due to certain features of its life cycle.     

Ecological factors influencing the composition of the parasite fauna of the European eel, Anguilla anguilla (L.), in Ireland

The parasites of 121 eels from three contrasting sites in the Corrib catchment area, western Ireland, were investigated. Thirteen species, Ergasilus gibbus, Diplostomum gasierostei, Diplostomum spathaceum, Sphaerostoma bramae. Bothrocephalus claviceps, Proteocephalus macrocephalus, Camallanus lacustris, Cucullanus truttae, Paraquimperia tenerrima, Raphidascaris acus, Acanthocephalus clavula, Acanthocephalus lucii and Pomphorhynchus laevis , were recorded. Two species, P. macrocephalus and P. tenerrima , have not previously been reported from Ireland. Microhabitat preferences of the parasites were noted. Variations in the occurrence and intensities of the parasites observed were analyzed in relation to sampling period, host habitat and characteristics of the eel populations studied. A variety of factors were shown to be of importance, including composition of the fish communities and distributional patterns of intermediate hosts and piscivorous birds. Differences were noted in the parasit-ocoenoses of eels in still and running water sites. The occurrence and intensities of infection of several parasite species were shown to be related to age and size of host: the occurrence of B. claviceps. C. truttae, P. tenerrima and R. acus was shown to be related to either age or size of eels, which is accounted for by the fact that eels become increasingly piscivorous with age and increasing size. Little evidence of interspecific interactions was noted.     

Species associations among larval helminths in an amphipod intermediate host

Larval helminths that share the same intermediate host may or may not also share the same definitive hosts. If one or more of these helminth species can manipulate the phenotype of the intermediate host, there can be great advantages or severe costs for other helminths resulting from co-occurring with a manipulator, depending on whether they have the same definitive host or not. Among 2372 specimens of the amphipod Echinogammarus stammeri collected from the river Brenta, northern Italy, there was a positive association between two acanthocephalan species with the same fish definitive hosts, the relatively common Pomphorhynchus laevis and the much less prevalent Acanthocephalus clavula.The number of cystacanths of P. laevis per infected amphipod, which ranged from one to five, did not influence the likelihood that the amphipod would also host A. clavula. A third acanthocephalan species, Polymorphus minutus,which matures in birds, showed no association with either of the two other species. These results show that associations among helminth species in intermediate hosts are not random, and are instead the product of selection favouring certain pathways of transmission.     

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.     

Role of the daily food ration of perch fry (Perca fluvialis) in infestation with Bunodera luciopercae (Trematoda, Allocreadiidae)

The list of intermediate hosts of Bunodera luciopercae is given. In Lake Syamozero they are represented by crustaceans Heterocope, Ophryoxus. Data on their role in the ration of juvenile perch are given. Diurnal dynamics of the ratio between infected and noninfected plankton in feeding and the process of the formation of fish infection have been studied.     

Proximate factors affecting the larval life history of Acanthocephalus lucii (Acanthocephala)

The growth and eventual size of larval helminths in their intermediate hosts presumably has a variety of fitness consequences. Therefore, elucidating the proximate factors affecting parasite development within intermediate hosts should provide insight into the evolution of parasite life histories. An experimental infection that resulted in heavy intensities of an acanthocephalan (Acanthocephalus lucii) in its isopod intermediate host (Asellus aquaticus) permitted the examination of parasite developmental responses to variable levels of resource availability and intraspecific competition. Isopods were infected by exposure to egg-containing fish feces, and larval infrapopulations were monitored throughout the course of A. lucii development. The relative rate of parasite growth slowed over time, and indications of resource constraints on developing parasites, e.g., crowding effects, were only observed in late infections. Consequently, the factors likely representative of resource availability to larval parasites (host size and molting rate) primarily affected parasite size in late infections. Moreover, at this stage of infection, competitive interactions, gauged by variation in worm size, seemed to be alleviated by greater resources, i.e., larger hosts that molted more frequently. The relatively rapid, unconstrained growth of young parasites may be worse for host viability than the slower, resource-limited growth of larger parasites.     

Utilization of brown trout Salmo trutta by Acanthocephalus clavula in an Irish lake: is this evidence of a host shift?

The population biology of the fish acanthocephalan Acanthocephalus clavula was described from 161 wild brown trout, Salmo trutta sampled over a two-year period in Clogher Lake in the west of Ireland. Overall prevalence of the parasite was 86% and the mean abundance was 53 worms per fish. Despite the presence of large numbers of worms in the trout very few females (2%) attained full reproductive maturity. This suggests that trout is an accidental host. A sample of yellow eels, Anguilla anguilla was examined at a different time from the same lake. The prevalence of A. clavula was 97% and the average abundance was 8 worms per fish. In contrast to the situation in trout, the proportion of female worms attaining full reproductive maturity was 61% fulfilling the expected characteristic of a preferred definitive host. The possible explanations for the very high abundance of A. clavula in trout are discussed and include the influence of fluctuations in host populations, host diet and the absence of a potential competitor.     

Quantitative effects of three species of parasites on a population of Three-spined sticklebacks, Gasterosteus aculeatus

The effects of Schistocephalus solidus, Diplostomum gasterostei and Echinorhynchus clavula on a population of Three-spined sticklebacks from a pond in Somerset were studied. Schistocephalus was present at the highest level of infection and had the greatest effects on the sticklebacks. It caused the fish to weigh less than uninfected ones of the same length, as shown by the lower condition factor, and to grow more slowly; and it delayed the sexual maturation of fish of both sexes and prevented many from breeding. Diplostomum also had a significant effect on the condition of the sticklebacks and all three parasite species probably caused death or predation of heavily infected fish. The quantitative effects of the parasites on the condition, length and weight of the fish were calculated; seasonal changes in the condition factor could then be seen, and the growth rates for both length and weight calculated.     

Diet and intestinal helminths in a population of perch, Perca flavescens

In 1967 and 1968 1310 Perca flavescens (Mitchill) were examined from Lake Opeongo, Ontario for changes in five diet categories and in incidence of eight intestinal helminths in relation to seasons, host size and sex. Male fish were less common than females and were smaller. A high proportion of fish (57.7 %) had empty stomachs, but of the four types of foods taken microcrustacea were high in spring, late summer and again in early winter; insects increased in summer with another peak in autumn; crayfish had a small peak in spring and a larger one in autumn; and fish were lowest in summer. Males ate more smaller foods (microcrustacea) than females, but females ate more larger foods (crayfish and fish); no sex difference was seen with regard to insects or empty stomachs. Male fish ate very few larger items, but females showed an increase in food size with increasing size of fish. There was no seasonality in incidence of fish free of all parasites, but two trematodes Bunodera sacculata Van Cleave & Mueller and B. luciopercae (Müller), and one cestode Proteocephalus pearsei La Rue showed seasonality with populations lowest in fish in summer; and one trematode Crepidostomum cooperi Hopkins and one nematode Dacnitoides cotylophora Ward & Magath showed seasonality with populations highest in summer. In addition one cestode Bothriocephalus sp., one nematode Spinitectus gracilis Ward & Magath and one acanthocephalan Leptorhynchoides thecatus (Linton) showed no seasonality, perhaps on account of the low incidence in the case of Bothriocephalus sp. and L. thecatus. No parasites were more common in males, but two (P. pearsei and L. thecatus) were more common in females. In both males and females, with increasing size fish, (i) fish free of all parasites decreased, but (ii) incidence of C. cooperi increased. Incidence of B. luciopercae, Bothriocephalus sp., S. gracilis and L. thecatus all increased with increasing size of fish only in females. The relationship between diet and incidence of intestinal helminths is discussed in the light of what is known of their life cycles. The exercise illustrates our ignorance of population ecology of fish parasites and the interactions of parasites and their host populations.     

Helminth and arthropod parasites of fishes in the mountain national parks of Canada

Sixteen parasite species including one monogenean, two digeneans, five cestodes, six nematodes. one acanthocephalan, and one copepod were collected from 257 fish of 13 species examined from 32 lake and nine stream sites in the mountain National Parks of British Columbia and Alberta. The following parasites are reported from Alberta for the first time: Bunodera luciopercae (Müller, 1776), Eubothrium rugosum Batsch 1786, Proteocephalus parallacticus MacLulich, 1943, Rhabdochona cascadilla Wigdor, 1918, Rhabdochona milleri Choquette, 1951, Contracaecum brachyurum (Ward & Magath, 1917), Bulbodacnitis truttae (Fabricius, 1794), Neochinorhynchus rutili (Mueller, 1780), and Ergasilus nerkae Roberts 1963. Tetraonchus variabilis Mizelle & Webb, 1953 and Rhabdochona milleri Choquette, 1951 are reported from British Columbia for the first time.     

The population biology of Camallanus lacustris (Zoega) in eels, Anguilla anguilla (Linnaeus), and their status as its host

The population biology of Camallanus lacustris (Zoega) and the status of the eel, Anguilla anguilla (Linnaeus), as its definitive host have been studied in a small Devon lake. No clear seasonal pattern in prevalence and abundance was observed, and recruitment of the new generation may occur in all seasons. However, reproduction exhibits some seasonality as adult nematodes were the commonest stage in the parasite population from early summer onwards and production of first-stage larvae appeared to occur primarily during this period. These larvae were viable, and shown experimentally to be infective to copepods. The occurrence of the nematode and its development to full maturity in eels, the absence of its typical definitive host ( Perca fluviatilis Linnaeus) from the lake and the absence of the nematode from other species of fish in the lake indicate that eels serve as the only, true definitive host of the nematode in this locality. It is suggested that account must be taken of this alternative host when the population of C. lacustris is investigated in the future.     

Selective predation on parasitized prey--a comparison between two helminth species with different life-history strategies.

In Lake Fjellfr?svatn, northern Norway, the larval helminths Cyathocephalus truncatus and Cystidicola farionis use Gammarus lacustris as intermediate hosts and Arctic charr (Salvelinus alpinus) as final hosts. There was sampled 1,433 live G. lacustris from the lake and 1,964 G. lacustris from stomach contents of the charr. Prevalence of infection were, respectively, 0.49% and 3.72% for C. truncatus, and 0.21% and 0.20% for C. farionis. Usually, only 1 parasite was present in each host, and the 2 parasite species never co-occurred. Gammarus lacustris amphipods parasitized by C. truncatus were positively selected by the Arctic charr and were consumed approximately 8 times as often as were the unparasitized amphipods or the amphipods infected with C. farionis. This suggests that G. lacustris amphipods infected with C. truncatus larvae are more susceptible to predation than noninfected specimens, probably because of parasite-induced alterations in behavior or visibility. Alternatively, this could also be explained by selection toward the largest G. lacustris specimens observed, which are also the most frequently parasitized amphipods. However, the data show clearly that this was not a result of size-selective predation by the charr. In contrast, the presence of C. farionis did not increase the susceptibility to predation of its intermediate host. The discrepancy between the 2 helminth species supports the hypothesis that parasite-increased susceptibility to predation is related to the life history strategies of the parasites.     

Effects of Acanthocephalus lucii (Acanthocephala) on intermediate host survival and growth: implications for exploitation strategies

Intermediate host exploitation by parasites is presumably constrained by the need to maintain host viability until transmission occurs. The relationship between parasitism and host survival, though, likely varies as the energetic requirements of parasites change during ontogeny. An experimental infection of an acanthocephalan (Acanthocephalus lucii) in its isopod intermediate host (Asellus aquaticus) was conducted to investigate host survival and growth throughout the course of parasite development. Individual isopods were infected by exposure to fish feces containing parasite eggs. Isopods exposed to A. lucii had reduced survival, but only early in the infection. Mean infection intensity was high relative to natural levels, but host mortality was not intensity dependent. Similarly, a group of naturally infected isopods harboring multiple cystacanths did not have lower survival than singly infected isopods. Isopods that were not exposed to the parasite exhibited sexual differences in survival and molting, but these patterns were reversed or absent in exposed isopods, possibly as a consequence of castration. Further, exposed isopods seemed to have accelerated molting relative to unexposed controls. Infection had no apparent effect on isopod growth. The effects of A. lucii on isopod survival and growth undermine common assumptions concerning parasite-induced host mortality and the resource constraints experienced by developing parasites.     

Observations on the occurrence of Allocreadium fasciatusi in Aplocheilus melastigma

Allocreadium fasciatusi is a common trematode occurring in the intestine of the fish Aplocheilus melastigma . The distribution of this fluke inside the fish host has been studied for a period of one year. The fluke occurred throughout the year, the incidence ranged from 20 to 88, and the number of parasites per infected host varied from 1 to 58. The fluke exhibited a seasonal cycle of occurrence with a peak both in incidence and intensity in the month of September. For the rest of the year the population remained constant and there was a balance between recruitment and loss from the trematode populations. The maturation cycle of A. fasciatusi also followed a seasonal pattern. Although the flukes were recruited in all the months, the peak invasion occurred in September. The large numbers of immature flukes received during this month became gravid by January. The availability of infective larvae was considered to be important in influencing the seasonal cycles.
The incidence and intensity of infection were found to be independent of the size of the fish but female fish were noted to be the more heavily infected.     

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.     

Perch (Perca fluviatilis L.) parasites reflect ecosystem conditions: a comparison of a natural lake and two acidic reservoirs in Finland

Parasite communities of perch were studied in a natural lake (Vetämäjärvi) and two reservoirs (the Kyrkösjärvi and Liikapuro reservoirs) located in Western Finland. All water bodies studied are small, shallow and humic. However, the reservoirs are more acidic than the lake (pH 5.9 and 5.3 vs. 6.4). Altogether, 18 parasite species were found, but the component communities were reduced in the harsh conditions of the reservoirs (12 and six species) as compared with the lake (17 species). In addition, the mean number of metazoan parasite specimens per fish was markedly lower in the reservoirs (64.3 and 14.3 specimens) than in the lake (116.1 specimens). Our prediction on the depauperation of parasite communities in reservoirs as compared with the lake was confirmed. Many ectoparasite groups were rare and most of the digenean parasites using molluscs as their intermediate host were absent from the reservoirs due to the inability of molluscs to tolerate the acidic water. The only digenean parasite found in the less acidic reservoir was Bunodera luciopercae. There were also species that preferred reservoirs as was expected. The protozoan ciliate Capriniana piscium benefiting acidic water and Acanthocephalus lucii, due to the abundance of its intermediate host, were most prevalent in the reservoir conditions.     

Life history and population biology of adult Acanthocephalus lucii (Acanthocephala: Echinorhynchidae)

The life history and population biology of adult A. lucii in perch, Perca fluviatilis L., from the Forth and Clyde canal, Scotland, was investigated during May 1979-September 1981. There was an annual cycle in the size of the parasite population; prevalence and abundance (+/- SE) were highest during late spring and summer (70-90% and 14 +/- 4.3 to 16 +/- 5.6 worms/fish, respectively) but declined during late autumn and reached a minimum during winter (50-60% and 2.1 +/- 0.9 to 3.2 +/- 0.6 worms/fish). Parasite maturation was associated with higher water temperatures during spring and summer and most shelled acanthors were probably produced during summer and fall. There was only 1 generation of A. lucii per year, although generations tended to overlap and individuals within each generation did not develop synchronously. The sex ratio of adults was initially near unity but favoured females in the later stages of the infection. The distribution of A. lucii among perch was highly aggregated and stomach content analysis suggested that this was partly due to heterogeneity in perch feeding behaviour. The negative binomial and Poisson lognormal models fitted the data on worm distribution. Seasonal changes in the degree of parasite aggregation were detected, but no conclusive evidence of density-dependent controls on parasite population growth was obtained.