The population biology of two species of eyefluke, Diplostomum spathaceum and Tylodelphys clavata, in roach and rudd

Infections of roach ( Rutilis rutilus ), and rudd ( Scardinius erythrophthalmus ), by the eyeflukes Diplostomun spathaceum and Tylodelphys clavata were studied in a lake over a three year period. T. clavata first appeared at the beginning of the study. Roach were heavily infected by both parasites, whereas the rudd contained very few specimens of either species. A small number of roach-rudd hybrids were also examined, and appeared to have a susceptibility intermediate between the parental types.
Levels of T. clavata in roach increased throughout the study. This parasite had a life span of a year or less in the fish, and was continuously lost from the population. This loss was balanced by a minor infection period in April/May and a major one extending from August until January. The fish continued to accumulate infections until they reached a size of about 130 mm, but thereafter intensity fluctuated about the level reached.
D. spathaceum infections similarly increased throughout the period of study, and there was a suggestion of a wave of infection in the spring. This parasite has a much longer life span, and levels of infection increased with fish size, only falling in the largest, and therefore oldest, hosts.
The major changes in the parasite populations in the roach were reflected, to a lesser extent, in the rudd. Both species were over-dispersed, but the data from roach was too heterogeneous to fit to any theoretical models, whereas that from the rudd was found to conform to a Negative Binomial distribution.     

Long term studies on the population biology of two specie of eyefluke, Diplostomum gasterostei and Tylodelphys clavata (Digenea: Diplostomatidae), concurrently infecting the eyes of perch, Perca fluviatilis

Changes in the size of the infrapopulations of metacercariae of Diplostomum gasterostei and Tylodelphys clavata were studied in a small lake for a period of seven years in perch Perca fluviatilis . Tylodelphys clavata appeared in the lake at the commencement of the study and its subsequent population increase and changes were typical of those of a species recently established in a new habitat. Population size increased slowly at first, then rapidly, then declined again as population density approached equilibrium level. By contrast, the population size of D. gasterostei declined throughout the study. Two processes were responsible for the decline: a decrease in the proportion of fish carrying heavy infections which was continuous throughout the seven years and a dramatic decline in recruitment rate in young fish during the last two years. In earlier years recruitment rate had changed annually and changes in overall infrapopulation size had largely reflected the recruitment rate two years previously. It was concluded that these two processes were independent of each other and related to changes in the population density of other species of eyeflukes. The decline in the proportion of heavily infected fish was significantly negatively correlated with the increase in population density of T. clavata and the decline in recruitment rate in young fish was coincident with the appearance and infrapopulation increase of T. podicipina metacercariae in the eyes of young perch. It is suggested that these two species interact negatively with D. gasterostei and are currently partitioning the perch habitat between themselves.     

The occurrence and life history of Asymphylodora kubanicum (Platyhelminthes: Digenea: Monorchidae) in the Worcester-Birmingham canal, with special reference to the feeding habits of the definitive host, Rutilus rutilus

Two hundred roach and over 400 molluscs were examined over a one year period to investigate the occurrence and life history of Asymphylodora kubanicum in the Worcester-Birmingham canal. Larval stages, infective to the fish definitive host, were present in molluscs throughout the year but did not show a seasonal fluctuation in numbers. Parasites in the intestine of the roach showed a marked annual cycle of occurrence and maturation: low winter infection levels preceded a dramatic increase in infection during the spring and summer. Maturation of the parasite population was rapid during the spring and summer and in late summer and early autumn the parasites laid their eggs and subsequently died. The death of parasites after egg-laying resulted in the low winter infection level during which time little recruitment occurred.
Roach became infected mainly in their third year when molluscs become a dominant component of their diet. Thereafter the older fishes (<2+) are all equally susceptible to infection, but the larger (older) fishes become more heavily infected because of the greater consumption of molluscs per fish. The sex of the fish made no difference to infection with the parasite.     

Long-term stability in the population levels of the eyefluke Tylodelphys podicipina (Digenea: Diplostomatidae) in perch

Annual changes in the population size of the metacercariae of Tylodelphys podicipina in the eyes of perch in a small eutrophic lake were studied over ten years by following changes in prevalence, abundance and overdispersion of parasites throughout the life of each year class of fish. The population increased rapidly in the first two years after its introduction to the lake, but for the next six years fluctuated within very narrowly constrained limits before declining as a result of a catastrophic decline in the perch population. No evidence of parasite-induced host mortality was found: the decline in parasite abundance throughout the life of each year class was due to density-independent parasite mortality within the fish as a result of natural variation in the life span of the metacercariae. Levels of infection in the first year of a year class were determined principally by transmission processes, as transmission could only occur over a very brief period when release of cercariae from parent generation snails coincided with the presence of perch fry. Transmission appeared to be density-independent, and infection levels in perch to be determined by the dimensions of the transmission' window' and subsequently modified by parasite mortality. No evidence of host responses or any other regulatory factor of fish origin was found, and the infrapopulations in perch appear to be non-equilibrial and unstable. Although the possibility of regulatory processes acting on infrapopulations of the parasite in other hosts remains, the importance of transmission windows in determining infrapopulation levels in fish is emphasized.     

The establishment and subsequent history of a population of Ligula intestinalis in roach Rutilis rutilis (L.)

The introduction, establishment and subsequent history of a population of Ligula intestinalis in roach, Rutilus rutilus , of a small lake was studied over a period of seven years. The introduction is believed to be a natural, chance colonization achieved by the return of Great Crested Grebes as permanent residents. Within two years the parasite had infected 33% of the roach between 60–120 mm in size. It exhibited a pronounced seasonal cycle in infection. Fish were often infected when only a few months old in autumn, and young fish and parasites over-wintered together. Parasite growth resumed in the following summer, when some further infections occurred. The majority of infected fish died in or before their second winter, possibly as a result of selective predation, but a small proportion survived into a third year. Dispersion of the parasite was close to random for most of the time. Multiple infections and slight over-dispersion occurred only in summer months. Infections were scarcer in large fish, due partly to selective mortality of infected fish and partly to a change in the host diet. The pattern of seasonal infection compared favourably with other localities, but prevalence and intensity of infection were consistently lower, parasite dispersion closer to random and multiple infections much scarcer than elsewhere. Infection levels and parasite growth rates varied from year to year, but these annual fluctuations were irregular and showed no consistent pattern. The Ligula population exhibited both regulatory and destabilizing features and there was no definite evidence that it was stable. It is suggested that Ligula is an opportunist species. It is endemic in some sites but in others, especially small lakes, the populations are unregulated and unstable and so persist for short periods only in a non-equilibrium state.     

Observations on the host-parasite relations of Thersitina gasterostei (Crustacea: Copepoda)

This study seeks to analyse the degree of infection of a euryhaline fish with the parasitic copepod Thersitina gasterostei in an extensive brackish-water system. For this purpose, a survey of the occurrence of the parasite on three-spined sticklebacks in the Norfolk Broads was conducted in Spring 1969. An attempt was made to correlate the incidence levels of T. gasterostei with variations in salinity. In addition, observations were made upon the location of the parasite on its host, the changes in maturation of the parasite during the sampling period and the differences in infection between three- and nine-spined sticklebacks.     

Seasonal dietary shifts enhance parasite transmission to lake salmonids during ice cover

Changes in abiotic and biotic factors between seasons in subarctic lake systems are often profound, potentially affecting the community structure and population dynamics of parasites over the annual cycle. However, few winter studies exist and interactions between fish hosts and their parasites are typically confined to snapshot studies restricted to the summer season whereas host‐parasite dynamics during the ice‐covered period rarely have been explored. The present study addresses seasonal patterns in the infections of intestinal parasites and their association with the diet of sympatric living Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) in Lake Takvatn, a subarctic lake in northern Norway. In total, 354 Arctic charr and 203 brown trout were sampled from the littoral habitat between June 2017 and May 2018. Six trophically transmitted intestinal parasite taxa were identified and quantified, and their seasonal variations were contrasted with dietary information from both stomachs and intestines of the fish. The winter period proved to be an important transmission window for parasites, with increased prevalence and intensity of amphipod‐transmitted parasites in Arctic charr and parasites transmitted through fish prey in brown trout. In Arctic charr, seasonal patterns in parasite infections resulted mainly from temporal changes in diet toward amphipods, whereas host body size and the utilization of fish prey were the main drivers in brown trout. The overall dynamics in the community structure of parasites chiefly mirrored the seasonal dietary shifts of their fish hosts.     

Interactions among co-infecting parasite species: a mechanism maintaining genetic variation in parasites?

Individuals of free-living organisms are often infected simultaneously by a community of parasites. If the co-infecting parasites interact, then this can add significantly to the diversity of host genotypexparasite genotype interactions. However, interactions between parasite species are usually not examined considering potential variation in interactions between different strain combinations of co-infecting parasites. Here, we examined the importance of interactions between strains of fish eye flukes Diplostomum spathaceum and Diplostomum gasterostei on their infectivity in naive fish hosts. We assessed the infection success of strains of both species in single-strain exposures and in co-exposures with a random strain of the other species. Parasite infection success did not consistently increase or decrease in the co-exposure treatment, but depended on the combinations of co-infecting parasite strains. This disrupted the relative infectivity of D. spathaceum strains observed in single-strain exposures. The infection success of D. gasterostei strains was independent of exposure type. These results suggest that interactions among parasite species may be strain specific and potentially promote maintenance of genetic polymorphism in parasite populations.     

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.     

The metazoan parasites of freshwater fishes in the Corrib catchment area, Ireland

A survey of the parasites of 243 fish belonging to 13 species, from the Corrib catchment area, western Ireland, was undertaken. At least 32 species of metazoan parasites were shown to be present. Diplostomum gasterostei, Tylodelphys clavata, Eubothrium salvelini, Cucullanus truttae, Raphidascaris acus, Ergasilus gibbus and Salmincola gordoni appear to be first records for Ireland and Phyllodistomum conostomum has not previously been reported from the British Isles. New host records involving Lamptera planeri, Salvelinus alpinus, Abramis brama, Noemacheilus barbatulus, Anguilla anguilla and Pungitius pungitius for the British Isles are given.     

Population dynamics of the philometrid nematode Margolisianum bulbosum in the southern flounder Paralichthys lethostigma (Pisces: Paralichthyidae) in South Carolina, USA

This is the first report of the philometrid nematode Margolisianum bulbosum Blaylock and Overstreet, 1999 from the southern flounder Paralichthys lethostigma on the east coast of the USA. Observation of adult female worms was used as an indication of the parasite's presence in the fish. Adult females were found only in P. lethostigma >50 mm total length. The overall prevalence was 74%, with a mean intensity of 5 female nematodes per parasitized fish. Infected flounders were found throughout the year with a statistically significant decrease in intensity in the winter months. Neither salinity, water temperature, fish gender nor fish age were found to influence either prevalence or intensity of infection in the flounder. While larvigerous (gravid) females were found throughout the year, the significant decrease in their occurrence during the summer through fall, in concert with an observed decrease in intensity of infection during the winter, indicated that the life cycle of this philometrid species is likely to be annual.     

The distribution and pathogenicity of larvae of Eustrongylides (Nematoda) in brown trout Salmo trutta L. in Fernworthy Reservoir, Devon

Infections of brown trout Salmo trutta L. by larvae of the nematode Eustrongylides sp. were studied over a period of two years in Fernworthy Reservoir, Devon. The parasite, here as elsewhere in Britain, is acquired only in the lake itself and not in the streams feeding it. Both native and introduced brown trout were infected but not rainbow trout which did not survive sufficiently long in the reservoir. The parasite showed a preference for female fish and for fish over 20 cm in length. Since these latter were approaching the end of their natural life span in the lake, there was insufficient time for the parasites to accumulate in older fish. No seasonal changes in infection levels or size composition of the parasite population were recorded. The larvae occurred in capsules, the majority of which were attached to the stomach wall. No local pathological effects associated with the capsules were observed and infested fish were of similar length, weight and condition to uninfected ones. The absence of harmful effects upon the trout in Fernworthy Reservoir is compared and contrasted with the effects of the parasite in other localities. It is concluded that although Eustrongylides can occasionally cause damage to some species of fish, there is no conclusive evidence that it does so to trout. but its unsightly appearance and harmful effects on birds renders it an undesirable parasite in any fishery.     

Reciprocal Interaction Matrix Reveals Complex Genetic and Dose-Dependent Specificity among Coinfecting Parasites

Abstract Understanding genetic specificity in factors determining the outcome of host-parasite interactions is especially important as it contributes to parasite epidemiology, virulence, and maintenance of genetic variation. Such specificity, however, is still generally poorly understood. We examined genetic specificity in interactions among coinfecting parasites. In natural populations, individual hosts are often simultaneously infected by multiple parasite species and genotypes that interact. Such interactions could maintain genetic variation in parasite populations if they are genetically specific so that the relative fitness of parasite genotypes varies across host individuals depending on (1) the presence/absence of coinfections and/or (2) the genetic composition of the coinfecting parasite community. We tested these predictions using clones of fish eye flukes Diplostomum pseudospathaceum and Diplostomum gasterostei. We found that interactions among parasites had a strong genetic basis and that this modified genetic variation in infection success of D. pseudospathaceum between single and multiple infections as well as across multiply infected host individuals depending on the genetic identity of the coinfecting D. gasterostei. The relative magnitude of these effects, however, depended on the exposure dose, suggesting that ecological factors can modify genetic interactions between parasites.     

Parasitism and Shoal Size in Juvenile Sticklebacks: Conflicting Selection Pressures from Different Ectoparasites?

In gregarious animals, group size correlates negatively with infection levels by some kinds of parasites and positively with infection by others. Conflicting selection pressures can be exerted simultaneously on a host species by different parasite species. Among stationary, mixed-species shoals of juvenile threespine sticklebacks, Gasterosteus aculeatus , and blackspotted sticklebacks, Gasterosteus wheatlandi , shoal size correlated differently with levels of infection by two species of ectoparasites. Stickleback shoal size correlated positively with infection levels by the copepod Thersitina gasterostei , which is transmitted among fish by short-lived planktonic larvae. In contrast, infection levels by the highly mobile crustacean parasite Argulus funduli did not decrease as shoal size increased, as predicted from an earlier laboratory experiment. The species composition of the different stickleback shoals also had an influence on some aspects of infection by these two parasite species. The contrasting mode of transmission of the two parasites results in one parasite species having a higher transmission rate among fish within large shoals, whereas the success of the other parasite species is independent of fish shoal size. The two ectoparasites may thus exert different selection pressures on stickleback shoal sizes.     

Accumulation of plerocercoids of Triaenophorus crassus in the second intermediate host Coregonus lavaretus and their effect on growth of the host

Patterns of accumulation of Triaenophorus crassus in its second intermediate host whitefish Coregonus lavaretus s.l. were studied between 1991 and 1996 from two host populations in two separate areas of Lake Saimaa, Finland. Whitefish were infected commonly with several T. crassus plerocercoids and the parasites were aggregated into the oldest hosts. In one host population the annual parasite accumulation was 0·9 parasites in all host age groups between 3 and 8 years. In the other host population the annual accumulation was 1·6 parasites in 3–5-year-old fish, but increased up to 3 to 4 parasites per year in fish over 5 years old. The increase did not coincide with the period of maturation or any increase in whitefish growth, both of which could alter the food intake of the fish. The sharp increase in the annual accumulation suggests a threshold intensity above which the probability of acquiring further parasites increases. In spite of a heavy aggregation of parasites there was no evidence of parasite-induced host mortality. The annual increase in mean abundance was not correlated with the mean annual weight increase in 2–4-year-old fish within cohorts. However, evidence of a negative effect of parasites on whitefish growth was revealed by back-calculation of lengths of uninfected and infected whitefish and correlations between length or weight of fish and intensity of infection with fish age. Both analyses suggested that larger young fish harboured more parasites than the smaller ones while in older fish the reverse was true, a pattern that has not been shown earlier for parasitized fish.     

Seasonal dynamics of the acanthocephalan Pomphorhynchm laevis (Müller, 1776) in its intermediate and preferred definitive hosts

No seasonal cycle was found in either the prevalence or the intensity of natural Pomphorhynchus laevis infections in Leuciscus cephalus . There was a slight seasonal change in female maturity distribution but only irregular fluctuations in the size structure of the adult parasite population throughout the year. Cystacanths were available in all seasons. Rates of parasite growth, maturation and mortality, but not establishment, increased with water temperature (or factors indirectly associated with elevated water temperature) in laboratory-infected Salmo gairdneri . Increased rates of parasite growth and maturation mask any marked shifts in the size and maturity structure of the adult parasite population which might otherwise be due to the higher turnover of adult parasites in the summer months.     

Host-parasite relationships of longnose dace, Rhinichthys cataractae, from the Ford River, Michigan.

A total of 1,115 longnose dace, Rhinichthys cataractae (family Cyprinidae), were examined for parasites from May 1983 through October 1986 from 3 localities in the Ford River in Michigan's Upper Peninsula. Thirteen parasite species (1 Monogenea, 2 Digenea, 2 Cestoda, 4 Nematoda, 1 Acanthocephala, 3 Protozoa) infected dace. The parasite faunas of dace, taxonomically and in species number, were similar between localities. Posthodiplostomum minimum minimum, Neascus sp., and Rhabdochona canadensis were the most common helminths infecting dace from each locality. The first 2 species did not exhibit consistent seasonal infection patterns between years, whereas the prevalence and mean intensity of R. canadensis in dace from the downriver locality were higher in summer 1983, 1984, and 1985. The intensity of infection of each of these helminth species significantly increased with host length. The prevalences and mean intensities of P. m. minimum, Neascus sp., and R. canadensis as well as the helminth infracommunity diversity were highest in dace from the upriver locality. The major factors that influenced parasite intensity were environmental factors that occurred when and where a fish began its life, the sequence of events that occurred in each habitat the fish encountered during its life, and the length of exposure (age of fish). Dace have isolationist helminth infracommunities arising from factors including ectothermy, a simple enteric system, restricted vagility, and being gape-limited. Allogenic helminths with indirect life cycles predominate in the depauperate helminth fauna of dace.     

Population and seasonal distribution of Procamallanus daccai in Eutropiichthys vacha in Kaptai Lake, Chittagong, Bangladesh

The population and seasonal distribution of Procamallanus daccai (Nematoda: Procamallanidae) in Eutropiichthys vacha (Siluridae) from the Kaptai Lake were studied for a period of 20 months during 1986-1987. A total of 208 fish were necropsied and 5652 worms were recovered, of which 2392 were from 87 infected male fish and 3260 were from 89 infected female fish. Prevalence values varied from 50% to 95%. The intensity and abundance of nematodes fluctuated over the study period and are statistically significant (F=14.87, P<0.0001), and showed a seasonal preference for the summer period (F=33.39, P<0.05). The mean intensity was the highest in the smallest size group at 66.3+/-12.7 which gradually decreased to 4+/-1.89 in the largest size group. No significant differences were found between parasites from male and female hosts, except in the summer. Male parasites were less abundant in the population. Female worms were grouped into immature, maturing and gravid and all these stages were present throughout the year. The highest occurrence of both immature and gravid P. daccai was found during the months of February to April each year. Peak recruitment seems to occur during this period due to the presence of high quantities of immature worms.     

Effects of water quality and trophic status on helminth infections in the cyprinid fish, Schizothorax niger Heckel, 1838 from three lakes in the Kashmir Himalayas

Water quality greatly influences the population density of aquatic biota, including parasites. In order to evaluate the relationship between fish parasites and water quality in Kashmir Himalayas, we assessed helminth parasite densities in Schizothorax niger Heckel, 1838 (an endemic cyprinid fish of Kashmir) from three lakes, namely Anchar, Manasbal and Dal, which reflected the varied stages of eutrophication. The overall prevalence of helminth infections was higher in the hypertrophic Anchar Lake (prevalence = 18.6%) compared to Manasbal Lake, which was the least eutrophied (prevalence = 6.4%). Furthermore, mean prevalence of monoxenous and heteroxenous parasites was higher in lakes containing higher levels of water degradation (Anchar and Dal). The mean number of helminth species per fish host was the highest in the hypertrophic lake (1.3 ± 0.3) in comparison to the least eutrophic lake (0.2 ± 1.5). Variability of calculated infection indices (prevalence, mean intensity and mean abundance) revealed that helminth parasite composition in the fish was affected by the lakes' environmental stress (degraded water quality). Therefore, data on the density of helminth parasites in fish can provide supplementary information on the pollution status of a water body.     

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.