Interspecific associations among larval helminths in fish

Various processes can generate associations between the larvae of different helminth species in their fish intermediate or paratenic host. We investigated the pairwise associations among larval helminth species in eight different fish populations, using two different coefficients of associations, in order to determine in what situations they are strongest. All helminth species included use the fish studied as either their second intermediate host or their paratenic host, and are acquired by the fish when it ingests an infected first intermediate host. The intensity of infection correlated positively with fish length for most helminth species. Pairs of species which both exhibited positive correlations with fish length tended to be more strongly associated with one another, although this tendency was not pronounced. Similarity in life cycle had a more important influence on pairwise associations. Among the 62 pairwise associations that could be computed, pairs of helminth species that shared both first intermediate hosts and definitive hosts were the most strongly associated, followed by pairs that shared only one other host, and finally by pairs that did not share other hosts. The results suggest that assemblages of larval helminth parasites in fish are not random collections of locally available species, but rather structured packets of larval parasites that travel together along common transmission routes.     

Evolution of complex life cycles in trophically transmitted helminths. I. Host incorporation and trophic ascent

Links between parasites and food webs are evolutionarily ancient but dynamic: life history theory provides insights into helminth complex life cycle origins. Most adult helminths benefit by sexual reproduction in vertebrates, often high up food chains, but direct infection is commonly constrained by a trophic vacuum between free‐living propagules and definitive hosts. Intermediate hosts fill this vacuum, facilitating transmission to definitive hosts. The central question concerns why sexual reproduction, and sometimes even larval growth, is suppressed in intermediate hosts, favouring growth arrest at larval maturity in intermediate hosts and reproductive suppression until transmission to definitive hosts? Increased longevity and higher growth in definitive hosts can generate selection for larger parasite body size and higher fecundity at sexual maturity. Life cycle length is increased by two evolutionary mechanisms, upward and downward incorporation, allowing simple (one‐host) cycles to become complex (multihost). In downward incorporation, an intermediate host is added below the definitive host: models suggest that downward incorporation probably evolves only after ecological or evolutionary perturbations create a trophic vacuum. In upward incorporation, a new definitive host is added above the original definitive host, which subsequently becomes an intermediate host, again maintained by the trophic vacuum: theory suggests that this is plausible even under constant ecological/evolutionary conditions. The final cycle is similar irrespective of its origin (upward or downward). Insights about host incorporation are best gained by linking comparative phylogenetic analyses (describing evolutionary history) with evolutionary models (examining selective forces). Ascent of host trophic levels and evolution of optimal host taxa ranges are discussed.     

Insight into the short-finned squid Illex coindetii (Cephalopoda: Ommastrephidae) feeding ecology: is there a link between helminth parasites and food composition?

Squids are especially frequent as paratenic hosts of helminth parasites, particularly to those that have elasmobranchs and mammals as final hosts. Among those parasite species, anisakid nematode larvae and cestode plerocercoids are most effectively transferred through the trophic chain by oegopsid squids. A total of 439 short-finned squids, Illex coindetii (245 males, 190 females and 4 unsexed) were sampled in the central part of the eastern Adriatic Sea in order to assess their helminth component community and parasite dynamics with respect to host sex, maturity, seasonality, and feeding behavior. Two larval helminths were isolated, i.e., larvae of Anisakis pegreffii, characterized by molecular tools at the species level, and plerocercoids of Phyllobothrium sp., with prevalences of 30.5% and 2.3%, respectively. Highly significant seasonal variation in diet consumption, congruent with seasonal variation in anisakid intensity, was observed, underlining the tight role of squid prey in the trophic transmission of parasite. Likewise, the highest helminth prevalence and intensity of infection was recorded in autumn, when the fish prey, mostly Maurolicus muelleri, comprised the greatest proportion of diet. This helped to assign the Adriatic broadtail shortfin squid not as a first, but as a second, paratenic host for the anisakid, unlike as suggested previously. The presence of larval A. pegreffii confirms its previously reported zoogeographical distribution in the Mediterranean and Adriatic Seas. The presence of 2 helminths in I. coindetii describes the feeding patterns of the squid, as well as clearly defined and coevolved predator-prey relationships.     

Manipulation of host-resource dynamics impacts transmission of trophic parasites

Many complex life cycle parasites rely on predator–prey interactions for transmission, whereby definitive hosts become infected via the consumption of an infected intermediate host. As such, these trophic parasites are embedded in the larger community food web. We postulated that exposure to infection and, hence, parasite transmission are inherently linked to host foraging ecology, and that perturbation of the host-resource dynamic will impact parasite transmission dynamics. We employed a field manipulation experiment in which natural populations of the eastern chipmunk (Tamias striatus) were provisioned with a readily available food resource in clumped or uniform spatial distributions. Using replicated longitudinal capture-mark-recapture techniques, replicated supplemented and unsupplemented control sites were monitored before and after treatment for changes in infection levels with three gastro-intestinal helminth parasites. We predicted that definitive hosts subject to food supplementation would experience lower rates of exposure to infective intermediate hosts, presumably because they shifted their diet away from the intermediate host towards the more readily available resource (sunflower seeds). As predicted, prevalence of infection by the trophically transmitted parasite decreased in response to supplemental food treatment, but no such change in infection prevalence was detected for the two directly transmitted parasites in the system. The fact that food supplementation only had an impact on the transmission of the trophically transmitted parasite, and not the directly transmitted parasites, supports our hypothesis that host foraging ecology directly affects exposure to parasites that rely on the ingestion of intermediate hosts for transmission. We concluded that the relative availability of different food resources has important consequences for the transmission of parasites and, more specifically, parasites that are embedded in the food web. The broader implications of these findings for food web dynamics and disease ecology are discussed.     

The trophic vacuum and the evolution of complex life cycles in trophically transmitted helminths

Parasitic worms (helminths) frequently have complex life cycles in which they are transmitted trophically between two or more successive hosts. Sexual reproduction often takes place in high trophic-level (TL) vertebrates, where parasites can grow to large sizes with high fecundity. Direct infection of high TL hosts, while advantageous, may be unachievable for parasites constrained to transmit trophically, because helminth propagules are unlikely to be ingested by large predators. Lack of niche overlap between propagule and definitive host (the trophic transmission vacuum) may explain the origin and/or maintenance of intermediate hosts, which overcome this transmission barrier. We show that nematodes infecting high TL definitive hosts tend to have more successive hosts in their life cycles. This relationship was modest, though, driven mainly by the minimum TL of hosts, suggesting that the shortest trophic chains leading to a host define the boundaries of the transmission vacuum. We also show that alternative modes of transmission, like host penetration, allow nematodes to reach high TLs without intermediate hosts. We suggest that widespread omnivory as well as parasite adaptations to increase transmission probably reduce, but do not eliminate, the barriers to the transmission of helminths through the food web.     

Factors influencing infection patterns of trophically transmitted parasites among a fish community: host diet,host–parasite compatibility or both?

Parasite infection patterns were compared with the occurrence of their intermediate hosts in the diet of nine sympatric fish species in a New Zealand lake. Stomach contents and infection levels of three gastrointestinal helminth species were examined from the entire fish community. The results highlighted some links between fish host diet and the flow of trophically transmitted helminths. Stomach contents indicated that all but one fish species were exposed to these helminths through their diet. Host feeding behaviour best explained infection patterns of the trematode Coitocaecum parvum among the fish community. Infection levels of the nematode Hedruris spinigera and the acanthocephalan Acanthocephalus galaxii, however, were not correlated with host diets. Host specificity is thus likely to modulate parasite infection patterns. The data indicate that host diet and host-parasite compatibility both contribute to the distribution of helminths in the fish community. Furthermore, the relative influence of encounter (trophic interactions between prey and predator hosts) and compatibility (host suitability) filters on infection levels appeared to vary between host-parasite species associations. Therefore, understanding parasite infection patterns and their potential impacts on fish communities requires determining the relative roles of encounter and compatibility filters within and across all potential host-parasite associations.     

Vulnerability and diet breadth predict larval and adult parasite diversity in fish of the Bothnian Bay

Recent studies of aquatic food webs show that parasite diversity is concentrated in nodes that likely favour transmission. Various aspects of parasite diversity have been observed to be correlated with the trophic level, size, diet breadth, and vulnerability to predation of hosts. However, no study has attempted to distinguish among all four correlates, which may have differential importance for trophically transmitted parasites occurring as larvae or adults. We searched for factors that best predict the diversity of larval and adult endoparasites in 4105 fish in 25 species studied over a three-year period in the Bothnian Bay, Finland. Local predator–prey relationships were determined from stomach contents, parasites, and published data in 8,229 fish in 31 species and in seals and piscivorous birds. Fish that consumed more species of prey had more diverse trophically transmitted adult parasites. Larval parasite diversity increased with the diversity of both prey and predators, but increases in predator diversity had a greater effect. Prey diversity was more strongly associated with the diversity of adult parasites than with that of larvae. The proportion of parasite species present as larvae in a host species was correlated with the diversity of its predators. There was a notable lack of association with the diversity of any parasite guild and fish length, trophic level, or trophic category. Thus, diversity is associated with different nodal properties in larval and adult parasites, and association strengths also differ, strongly reflecting the life cycles of parasites and the food chains they follow to complete transmission.     

The evolution of complex life cycles when parasite mortality is size- or time-dependent

In complex cycles, helminth larvae in their intermediate hosts typically grow to a fixed size. We define this cessation of growth before transmission to the next host as growth arrest at larval maturity (GALM). Where the larval parasite controls its own growth in the intermediate host, in order that growth eventually arrests, some form of size- or time-dependent increase in its death rate must apply. In contrast, the switch from growth to sexual reproduction in the definitive host can be regulated by constant (time-independent) mortality as in standard life history theory. We here develop a step-wise model for the evolution of complex helminth life cycles through trophic transmission, based on the approach of Parker et al. [2003a. Evolution of complex life cycles in helminth parasites. Nature London 425, 480-484], but which includes size- or time-dependent increase in mortality rate. We assume that the growing larval parasite has two components to its death rate: (i) a constant, size- or time-independent component, and (ii) a component that increases with size or time in the intermediate host. When growth stops at larval maturity, there is a discontinuous change in mortality to a constant (time-independent) rate. This model generates the same optimal size for the parasite larva at GALM in the intermediate host whether the evolutionary approach to the complex life cycle is by adding a new host above the original definitive host (upward incorporation), or below the original definitive host (downward incorporation). We discuss some unexplored problems for cases where complex life cycles evolve through trophic transmission.     

Host sharing and host manipulation by larval helminths in shore crabs: cooperation or conflict?

Larval helminths of different species that share the same intermediate host and are transmitted by predation to the same definitive host may cooperate in their attempts to manipulate the behaviour of the intermediate host, while at the same time having conflicts of interests over the use of host resources. A few studies have indicated that intermediate hosts harbouring larval helminths have altered concentrations of neurotransmitters in their nervous system, and thus measuring levels of neurotransmitters in host brains could serve to assess the respective and combined effect of different helminth species on host behaviour. Here, we investigate potential cooperation and conflict among three helminths in two species of crab intermediate hosts. The acanthocephalan Profilicollis spp., the trematode Maritrema sp. and an acuariid nematode, all use Macrophthalmus hirtipes (Ocypodidae) as intermediate host, whereas Profilicollis and Maritrema also use Hemigrapsus crenulatus (Grapsidae). All three helminths mature inside gulls or other shore birds. There was a significant decrease in the mean volume of Profilicollis cystacanths as the intensity of infection by this parasite increased in H. crenulatus, the only host in which this was investigated; however, there was no measurable effect of other helminth species on the size of acanthocephalans, suggesting no interspecific conflict over resource use within crabs. There was, in contrast, evidence of a positive interspecific association between the two most common helminth species: numbers of Profilicollis and Maritrema were positively correlated among crabs, independently of crab size, in M. hirtipes but not H. crenulatus. More importantly, we found that the total number of larval helminths per crab correlated significantly, and negatively, with concentrations of serotonin in crab brains, again only in M. hirtipes; numbers of each parasite species separately did not covary in either crab species with serotonin or dopamine, the other neurotransmitter investigated in this study. The relationship with serotonin appears due mainly to numbers of Profilicollis and Maritrema and not to nematodes. This is the first demonstration of a potentially synergistic manipulation of host behaviour by different helminth species, one that appears host-specific; our results also point toward the neurobiological mechanism underlying this phenomenon.     

Intra- and interspecific competition among helminth parasites: effects on Coitocaecum parvum life history strategy, size and fecundity

Larval helminths often share intermediate hosts with other individuals of the same or different species. Competition for resources and/or conflicts over transmission routes are likely to influence both the association patterns between species and the life history strategies of each individual. Parasites sharing common intermediate hosts may have evolved ways to avoid or associate with other species depending on their definitive host. If not, individual parasites could develop alternative life history strategies in response to association with particular species. Three sympatric species of helminths exploit the amphipod Paracalliope fluviatilis as an intermediate host in New Zealand: the acanthocephalan Acanthocephalus galaxii, the trematode Microphallus sp. and the progenetic trematode Coitocaecum parvum. Adult A. galaxii and C. parvum are both fish parasites whereas Microphallus sp. infects birds. We found no association, either positive or negative, among the three parasite species. The effects of intra- and interspecific interactions were also measured in the trematode C. parvum. Both intra- and interspecific competition seemed to affect both the life history strategy and the size and fecundity of C. parvum. Firstly, the proportion of progenesis was higher in metacercariae sharing their host with Microphallus sp., the bird parasite, than in any other situation. Second, the intensity of intraspecific competition apparently constrained the ability of metacercariae to adopt progenesis and limited both the growth and egg production of progenetic individuals. These results show that the life history strategy adopted by a parasite may be influenced by other parasites sharing the same host.     

Helminth communities of native and introduced fishes in Lake Pátzcuaro, Michoacán, México

In Lake Pátzcuaro in the Mesa Central of México, a total of 19 species of helminths was found in 598 fishes and comprised five digeneans, two monogeneans, four cestodes, one acanthocephalan and seven nematodes, of which ten species were represented by larval or immature states. The richest and most diverse helminth communities were found in the native carnivorous goodeid Alloophorus robustus. In general, the helminth communities in the different fish species were not particularly species rich and the parasite assemblages were numerically dominated by larvae of the bird trematode, Posthodiplostomum minimum. Patterns of helminth community richness and diversity were similar to those previously observed in north-temperate freshwater fishes. Most enteric helminths occurred with low abundance and only a small proportion of the gut helminth communities was numerically dominated by any one species. Helminths dominating their enteric communities showed some level of host specificity. Helminth communities in carnivorous fish species were generally richer than those in herbivores and detritivores, with the exception of the predominantly herbivorous Goodea atripinnis. The helminth fauna of introduced fishes, Cyprinus carpio, Micropterus salmoides and Oreochromis niloticus , consisted of either few or no host-specific adult helminth(s) translocated from their original geographical areas and by larval stages of helminths of piscivorous birds. Based on the geological history of the area and the biogeography of the endemic fish fauna, it is hypothesized that host-switching and relationships with the nearctic fauna have been fundamental in determining the helminth fauna of the endemic fish hosts.     

Larval helminths in intermediate hosts: does competition early in life determine the fitness of adult parasites?

Density-dependent effects on parasite fitness have been documented from adult helminths in their definitive hosts. There have, however, been no studies on the cost of sharing an intermediate host with other parasites in terms of reduced adult parasite fecundity. Even if larval parasites suffer a reduction in size, caused by crowding, virtually nothing is known about longer-lasting effects after transmission to the definitive host. This study is the first to use in vitro cultivation with feeding of adult trematodes to investigate how numbers of parasites in the intermediate host affect the size and fecundity of adult parasites. For this purpose, we examined two different infracommunities of parasites in crustacean hosts. Firstly, we used experimental infections of Maritrema novaezealandensis in the amphipod, Paracalliope novizealandiae, to investigate potential density-dependent effects in single-species infections. Secondly, we used the crab, Macrophthalmus hirtipes (Ocypodidae), naturally infected by the trematodes, M. novaezealandensis and Levinseniella sp., the acanthocephalan, Profilicollis spp., and an acuariid nematode. These four helminths all develop and grow in their crustacean host before transmission to their bird definitive host by predation. In experimental infections, we found an intensity-dependent establishment success, with a decrease in the success rate of cercariae developing into infective metacercariae with an increasing dose of cercariae applied to each amphipod. In natural infections, we found that M. novaezealandensis-metacercariae achieved a smaller volume, on average, when infrapopulations of this parasite were large. Small metacercariae produced small in vitro-adult worms, which in turn produced fewer eggs. Crowding effects in the intermediate host thus were expressed at the adult stage in spite of the worms being cultured in a nutrient-rich medium. Furthermore, excystment success and egg-production in M. novaezealandensis in naturally infected crabs were influenced by the number of co-occurring Profilicollis cystacanths, indicating interspecific interactions between the two species. Our results thus indicate that the infracommunity of larval helminths in their intermediate host is interactive and that any density-dependent effect in the intermediate host may have lasting effects on individual parasite fitness.     

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

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

Metazoan parasites of African annual killifish (Nothobranchiidae): abundance,diversity, and their environmental correlates

Estimates of biodiversity and its global patterns are affected by parasite richness and specificity. Despite this, parasite communities are largely neglected in biodiversity estimates, especially in the tropics. We studied the parasites of annual killifish of the genus Nothobranchius that inhabit annually desiccating pools across the African savannah and survive the dry period as developmentally arrested embryos. Their discontinuous, non‐overlapping generations make them a unique organism in which to study natural parasite fauna. We investigated the relationship between global (climate and altitude) and local (pool size, vegetation, host density and diversity, and diversity of potential intermediate hosts) environmental factors and the community structure of killifish parasites. We examined metazoan parasites from 21 populations of four host species (Nothobranchius orthonotus, N. furzeri, N. kadleci, and N. pienaari) across a gradient of aridity in Mozambique. Seventeen parasite taxa were recorded, with trematode larval stages (metacercariae) being the most abundant taxa. The parasites recorded were both allogenic (life cycle includes non‐aquatic host; predominantly trematodes) and autogenic (cycling only in aquatic hosts; nematodes). The parasite abundance was highest in climatic regions with intermediate aridity, while parasite diversity was associated with local environmental characteristics and positively correlated with fish species diversity and the amount of aquatic vegetation. Our results suggest that parasite communities of sympatric Nothobranchius species are similar and dominated by the larval stages of generalist parasites. Therefore, Nothobranchius serve as important intermediate or paratenic hosts of parasites, with piscivorous birds and predatory fish being their most likely definitive hosts.     

A gastropod scavenger serving as paratenic host for larval helminth communities in shore crabs

The whelk Cominella glandiformis is an important predator-scavenger of New Zealand intertidal ecosystems; a few whelks can quickly eat all the soft tissues of recently dead crabs. In this study, we demonstrate that whelks can also ingest and act as paratenic hosts for at least 4 helminth species that use crabs as intermediate hosts: metacercariae of the trematode Maritrema sp. and of another unidentified trematode, larval acuariid nematodes, and cystacanths of the acanthocephalans Profilicollis spp. Large whelks ingest disproportionately more helminth larvae than small whelks, but the survival of parasites during their short stay in the whelks is not affected by whelk size. The majority of metacercariae and nematodes are passed out in whelk feces within 3 days of ingestion, whereas the few cystacanths found did not leave whelks until after that time; no parasite was left in whelks 5 days postingestion. Survival of all 4 helminth species was generally very high, though it decreased day by day in 2 species. Given that the avian definitive hosts of all 4 helminths also eat whelks, our results indicate that alternative transmission pathways exist and that parasites can take routes through food webs that are too often ignored.     

How to catch a parasite: Parasite Niche Modeler (PaNic) meets Fishbase

Parasite Niche Modeler (PaNic) is a free online software tool that suggests potential hosts for fish parasites. For a particular parasite species from the major helminth groups (Acanthocephala, Cestoda, Monogenea, Nematoda, Trematoda), PaNic takes data from known hosts (maximum body length, growth rate, life span, age at first maturity, trophic level, phylogeny, and biogeography) and hypothesizes similar fish species that might serve as hosts to that parasite. Users can give varying weights to host attributes and create custom models. In addition to suggesting plausible hosts (with varying degrees of confidence), the models indicate known host species that appear to be outliers in comparison to other known hosts. These unique features make PaNic an innovative tool for addressing both theoretical and applied questions in fish parasitology. PaNic can be accessed at < http://purl.oclc.org/fishpest >.     

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.     

The role of the rock cod Notothenia coriiceps Richardson, 1844 in the life-cycle of Antarctic parasites

Fifty specimens of Notothenia coriiceps caught in Potter Cove, King George Island, were examined for ecto- and endoparasites. Of the 22 parasite species found, 18 were helminths, 2 were hirudineans and 2 were crustaceans. The isopod Aega antarctica and an unidentified hirudinean are reported for the first time from this fish host. Dominant parasites were the adults of Aspersentis megarhynchus, the invasive stage of Corynosoma spp. (cystacanth) and the adults of Macvicaria pennelli, with respective prevalences of infestation of 94, 76 and 74%. The preferred sites of infestation were the pylorus and intestine, where five different larval (nematodes and cestodes) and eight adult (digeneans and acanthocephalans) parasite species were found. No adult nematodes and cestodes were found and no parasites could be isolated from the musculature. The results of the present study are related to previous findings on the parasite fauna of N. coriiceps. The comparison implies a high parasite diversity in this benthic Antarctic fish species. Most parasites found appear to have a wide range of distribution within Antarctic waters together with a low host specificity. Besides its role as final host for several species of trematodes and acanthocephalans, N. coriiceps serves as transmitter of parasite larvae to piscivorous birds and seals. It is concluded that the parasite fauna in Antarctic fish species provides important insights into the different habitat use and trophic relationship of their fish hosts. Received: 11 September 1997 / Accepted: 12 January 1998     

Helminth communities of the lesser sandeel Ammodytes tobianus L. off the west coast of Ireland

The helminth parasites present in 412 lesser sandeels (Ammodytes tobianus) taken from June 1996 to May 1997 from the Aran Islands on the west coast of Ireland were examined. Ten helminth parasite species were recorded, and more than 92% of the sandeels were infected with at least 1 helminth species. Seven of the species were digeneans, including Brachyphallus crenatus, Hemiurus communis, Derogenes varicus, Lecithaster gibbosus, Opechona bacillaris, Cryptocotyle lingua, and Galactosomum lacteum; 2 nematodes, including Hysterothylacium sp. and Contracaecum sp.; and 1 cestode, Scolex pleuronectis. Three of the 7 digenean species were either larvae or immature. Only 2 species, the digeneans G. lacteum and H. communis, had prevalences greater than 50%. The dominant species was G. lacteum, accounting for 67% of all parasites present. The relationship between spawned groups, host length, and season versus the abundance, prevalence, species richness, and the total number of parasites in the infracommunities was investigated. No difference was found between the parasite communities of the 2 spawning races of the host population. Mean abundance and prevalence of the different parasite species showed seasonal variation. Numbers of parasite species and numbers of parasites increased with fish length. The role of A. tobianus as an intermediate host for helminths was assessed; it was determined that most were infectious to birds or mammals, with the majority of the parasite species being autogenic (infectious to fish). The mean number of parasites per fish was nearly a quarter of the value recorded for A. tobianus in the North Sea, where a much higher intensity of infection was recorded.     

Methylmercury levels in a parasite (<Emphasis Type="Italic">Apophallus brevis</Emphasis> metacercariae) and its host,yellow perch (<Emphasis Type="Italic">Perca flavescens</Emphasis>)

The biomagnification of methylmercury (MeHg) amongst trophic levels results in high levels of this compound in many freshwater fish species. The role of parasites in MeHg cycling and trophic transfer in freshwater systems is largely unknown. This study examined the potential for metacercariae of Apophallus brevis to accumulate and biomagnify MeHg from their second intermediate host, yellow perch, Perca flavescens. Contrary to our prediction that MeHg levels would be higher in parasites than in the host muscle tissue in which they are embedded, we found that concentrations were similar. The lack of increase in MeHg levels from host to parasite may be due to limited assimilation of host muscle tissue or, in part, to low parasite metabolism. Parasite load did not reduce fish growth and subsequently alter MeHg concentrations. This study suggests that relationships between larval parasites and their hosts do not conform to typical patterns of MeHg biomagnification seen in aquatic systems.