Effects of Herbicides and the Chytrid Fungus Batrachochytrium dendrobatidis on the growth, development and survival of Larval American Toads (Anaxyrus americanus).

Pesticides and pathogens adversely affect amphibian health, but their interactive effects are not well known. We assessed independent and combined effects of two agricultural herbicides and the fungal pathogen Batrachochytrium dendrobatidis (Bd) on the growth, development and survival of larval American toads (Anaxyrus americanus). Wild-caught tadpoles were exposed to four concentrations of atrazine (0.18, 1.8, 18.0, 180 µg/L) or glyphosate (7, 70, 700, 7000 µg a.e./L), respectively contained in Aatrex® Liquid 480 (Syngenta) or Vision® Silviculture Herbicide (Monsanto) for 14 days, followed by two doses of Bd. At day 14, atrazine had not affected survival, but it non-monotonically affected growth. Exposure to the highest concentration of glyphosate caused 100% mortality within 4 days, while lower doses had an increasing monotonic effect on growth. At day 65, tadpole survival was unaffected by atrazine and the lower doses of glyphosate. Neither herbicide demonstrated an interaction effect with Bd on survival, but exposure to Bd increased survival among both herbicide-exposed and herbicide-control tadpoles. At day 60, tadpoles exposed to the highest concentration of atrazine remained smaller than controls, indicating longer-term effects of atrazine on growth, but effects of glyphosate on growth disappeared. Growth was unaffected by any herbicide-fungal interaction but was positively affected by exposure to Bd following exposure to atrazine. Atrazine exhibited a slowing and non-monotonic effect on Gosner developmental stage, while exposure to Bd tended to speed up development and act antagonistically toward the observed effect of atrazine. Overall, atrazine, glyphosate and Bd all showed a potential to modulate larval toad growth and development.

Population growth of Gyrodactylus kobayashii in goldfish (Carassius auratus) associated with host density.

Host density is a key regulatory factor in parasite transmission. The goldfish (Carassius auratus)-Gyrodactylus kobayashii model was used to investigate effects of host density on population growth of gyrodactylids. A donor fish infected by five gravid gyrodactylids was mixed with 11 parasite-free goldfish at five host densities. There was a significant positive correlation between host density and mean abundance of G. kobayashii throughout the 58-day experiment. During early infection (days 15­24), mean abundance in medium high (0.5 fish L−1) and high host density groups (1 and 2 fish L−1) was significantly higher than that in the low host density groups (0.125 and 0.25 fish L−1). At high host density, prevalence increased more rapidly, and the peak prevalence was higher. Fitting of an exponential growth model showed that the population growth rate of the parasite increased with host density. A hypothesis was proposed that higher host density contributed to increased reinfection of detached gyrodactylids. A reinfection experiment was designed to test this hypothesis. Both mean abundance and prevalence at a host density of 1 fish L−1 were significantly higher than those at 0.25 fish L−1 on days 1 and 3, which suggested that more reinfections of G. kobyashii occurred at the higher host density. Density-dependent transmission during the early infection was an important contributor of population growth of G. kobayashii, as well as density-dependent reinfection of the detached gyrodactylids.

Effects of multiple stressors on northern leopard frogs in agricultural wetlands.

Natural and anthropogenic stressors, including parasites and pesticides, may induce oxidative stress in animals. Measuring oxidative stress responses in sentinel species that are particularly responsive to environmental perturbations not only provides insight into host physiology but is also a useful readout of ecosystem health. Newly metamorphosed northern leopard frogs (Lithobates pipiens), a sentinel species, were collected from agricultural and non-agricultural wetlands exposed to varying concentrations of the herbicide atrazine. Significant effects of certain parasites' abundance and their interaction with atrazine exposure on frog oxidative stress were identified. Specifically, increased protein levels were detected in frogs infected with echinostome metacercariae. In addition, the nematode Oswaldocruzia sp. was significantly associated with increased thiol concentration and catalase activity. Significant parasite × atrazine interactions were observed for atrazine exposure and the abundance of Oswaldocruzia sp. on thiol, as thiol concentrations increased with parasite abundance at low atrazine localities and decreased in high atrazine wetlands. In addition, a significant interaction between the abundances of Oswaldocruzia sp. and gorgoderid trematodes on thiol concentrations was observed. These findings demonstrate that studies of oxidative stress on animals in natural ecosystems should account for the confounding effects of parasitism, particularly for amphibians in agricultural landscapes.

Effects of goldfish (Carassius auratus) population size and body condition on the transmission of Gyrodactylus kobayashii (Monogenea).

Field surveys indicate that host population size, rather than density, is the most important determinant of monogenean infection dynamics. To verify this prediction, epidemic parameters were monitored for 70 days at five host population sizes held at constant density using a goldfish - Gyrodactylus kobayashii laboratory model. During the first 20 days, the rate of increase of prevalence and mean abundance was faster in small host populations. Total mean prevalence and total mean abundance throughout the experiment were not significantly affected by host population sizes. Higher transmission rates were detected in larger host populations. However, there were no significant differences in effective contact rates among the five host populations on each sampling day during the first 20 days, implying that contact rates may be saturated at a sufficiently high host density. These results demonstrate that the epidemic occurs more quickly in smaller host populations at the beginning of the experiment. However, the epidemic is independent of the host population size due to the similar effective contact rates in the five population sizes. Significant negative influence of the initial body condition (Kn) of uninfected goldfish on total mean abundance of parasites suggests that susceptibility of hosts is also a determinant of parasite transmission.

Three-spined stickleback Gasterosteus aculeatus, as a possible paratenic host for salmonid nematodes in a subarctic lake.

In Takvatn, a subarctic lake in northern Norway, 35 of 162 three-spined sticklebacks examined were infected with 106 specimens of third-stage larvae of Philonema oncorhynchi. The prevalence and mean intensity of P. oncorhynchi were 10 % and 2.0 in 2013 and 24 % and 3.0 in 2014, respectively. A single specimen of Cystidicola farionis was found in an additional sample. While the latter is considered an accidental infection, three-spined sticklebacks may function as paratenic hosts of P. oncorhynchi, potentially enhancing its transmission to salmonids due to their central role in the lacustrine food web of this subarctic lake.

Parasites as biological tags of marine, freshwater and anadromous fishes in North America from the Tropics to the Arctic.

Parasites have been considered as natural biological tags of marine fish populations in North America for almost 75 years. In the Northwest Atlantic, the most studied species include Atlantic cod (Gadus morhua), Atlantic herring (Clupea harengus) and the redfishes (Sebastes spp.). In the North Pacific, research has centred primarily on salmonids (Oncorhynchus spp.). However, parasites have been applied as tags for numerous other pelagic and demersal species on both the Atlantic and Pacific coasts. Relatively few studies have been undertaken in the Arctic, and these were designed to discriminate anadromous and resident salmonids (Salvelinus spp.). Although rarely applied in fresh waters, parasites have been used to delineate certain fish stocks within the Great Lakes-St Lawrence River basin. Anisakid nematodes and the copepod Sphyrion lumpi frequently prove useful indicators in the Northwest Atlantic, while myxozoan parasites prove very effective on the coast and open seas of the Pacific Ocean. Relative differences in the ability of parasites to discriminate between fish stocks on the Pacific and Atlantic coasts may be due to oceanographic and bathymetric differences between regions. Molecular techniques used to differentiate populations and species of parasites show promise in future applications in the field.

Parasite communities of two three-spined stickleback populations in subarctic Norway--effects of a small spatial-scale host introduction.

Co-introduction and colonization of parasites with the introduction of new host species into aquatic habitats may depend on the host specificity and dispersal capabilities of the parasites. We compared the metazoan parasite community of an introduced three-spined stickleback (Gasterosteus aculeatus) population with that of the nearby source population in subarctic Norway. As expected from a small spatial scale (5 km), the parasite component communities in the two lakes were highly similar. All identifiable allogenic parasite taxa (Diphyllobothrium dendriticum, Diphyllobothrium ditremum, Diphyllobothrium spp., Schistocephalus solidus, Apatemon sp. and Diplostomum spp.) were also observed in both lakes while inter-lake differences were driven by autogenic parasite taxa (Eubothrium spp., Crepidostomum spp., Nematoda spp., Proteocephalus sp. and Gyrodactylus arcuatus). Contrary to expectation, the total number of parasite taxa was higher in the introduced stickleback population (12) compared to that found in the source population (9) with three parasite taxa (Eubothrium spp., Crepidostomum spp., Nematoda spp.) only occurring in the introduced population. These parasites were uncommon however and normally restricted to salmonids. Sticklebacks from both populations were heavily infected, particularly with eye-infecting metacercariae. Sequences from the DNA barcode region of cytochrome oxidase 1 indicated that these include Diplostomum lineage 6, a member of the Diplostomum baeri complex and a member of the Strigeinae. Despite high similarity between the two component communities, quantitative inter-lake differences were found at the infracommunity level. At this scale, parasite intensity was significantly higher in the source population for the two autogenic stickleback specialists: G. arcuatus and Proteocephalus sp., assumed to be the autogenic stickleback specialist Proteocephalus filicollis. Parasite infracommunities within each lake also resembled each other significantly more than infracommunities between lakes, primarily driven by the allogenic cestode D. ditremum, as well as G. arcuatus and Proteocephalus sp. Overall, quantitative dissimilarities between the two parasite communities were possibly explained by inter-lake differences in the density of sticklebacks and intermediate hosts.

Effects of a major municipal effluent on the St. Lawrence River: A case study.

The St. Lawrence River (SLR) is the second largest waterway in North America. The discharge of the City of Montreal wastewater treatment plant (WWTP) represents the largest volume of treated wastewaters being released into the river. It also ranks as the largest sewage treatment plant of its kind in North America. Over the last decade, intensive multidisciplinary research has focused on assessing the impacts of Montreal wastewater effluents on the SLR. We describe the major findings of these investigations, including the determination of the fate of contaminants, bioaccumulation in fish and invertebrates, ecotoxicological measurements of aquatic animal health, evaluation of endocrine disruption, parasitism in fish, and combined effects of multiple stressors on the SLR. Impacts of the effluents from the WWTP on aquatic organisms from the SLR are both toxicological and ecological, demonstrating the need for an integrated view of the impacts of municipal effluents on aquatic ecosystems.

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.

Altered trophic pathway and parasitism in a native predator (Lepomis gibbosus) feeding on introduced prey (Dreissena polymorpha).

Populations of invasive species tend to have fewer parasites in their introduced ranges than in their native ranges and are also thought to have fewer parasites than native prey. This 'release' from parasites has unstudied implications for native predators feeding on exotic prey. In particular, shifts from native to exotic prey should reduce levels of trophically transmitted parasites. We tested this hypothesis in native populations of pumpkinseed sunfish (Lepomis gibbosus) in Lake Opinicon, where fish stomach contents were studied intensively in the 1970s, prior to the appearance of exotic zebra mussels (Dreissena polymorpha) in the mid-1990s. Zebra mussels were common in stomachs of present-day pumpkinseeds, and stable isotopes of carbon and nitrogen confirmed their importance in long-term diets. Because historical parasite data were not available in Lake Opinicon, we also surveyed stomach contents and parasites in pumpkinseed in both Lake Opinicon and an ecologically similar, neighboring lake where zebra mussels were absent. Stomach contents of pumpkinseed in the companion lake did not differ from those of pre-invasion fish from Lake Opinicon. The companion lake, therefore, served as a surrogate "pre-invasion" reference to assess effects of zebra mussel consumption on parasites in pumpkinseed. Trophically transmitted parasites were less species-rich and abundant in Lake Opinicon, where fish fed on zebra mussels, although factors other than zebra mussel consumption may contribute to these differences. Predation on zebra mussels has clearly contributed to a novel trophic coupling between littoral and pelagic food webs in Lake Opinicon.

Waterborne zinc alters temporal dynamics of guppy Poecilia reticulata epidermal response to Gyrodactylus turnbulli (Monogenea).

The present study assessed the histological changes in the epidermis of Poecilia reticulata induced by the combined effects of an ectoparasite Gyrodactylus turnbulli and differing concentrations of waterborne zinc (Zn). Infected guppies were exposed to 0, 15, 30, 60, or 120 µg Zn l-1 and monitored over 3 wk during the exponential increase in parasite numbers on the fish. The fish epidermis responded within 3 d to G. turnbulli infection with a rapid increase in epidermal thickness and a modest increase in number, but not size or composition, of mucous cells. In contrast, in the presence of combined waterborne Zn and infection, mucous cell numbers declined rapidly. As the parasite numbers increased, the epidermis remained thicker than normal, and the number and size of mucous cells decreased. The addition of Zn led to a dramatic thickening of the epidermis during the exponential growth of the parasite population. Mucous cell numbers remained depressed. Temporal changes in mucous cell size were Zn concentration dependent. At 60 µg Zn l-1, cells returned to normal size as infection progressed, whereas they remained extremely small at 120 µg Zn l-1. Changes in mucin composition previously reported in response to Zn alone were subdued in the presence of the parasite except at 60 µg Zn l-1, where all cells contained only acidic mucins. Together these results demonstrate that, on exposure to both Zn and G. turnbulli infection, the epidermal response is initially a protective response to both stressors, and then mainly driven by the increased parasite burden.

Effects of herbicides and the chytrid fungus Batrachochytrium dendrobatidis on the health of post-metamorphic northern leopard frogs (Lithobates pipiens).

Effects of exposure to contaminants such as pesticides along with exposure to pathogens have been listed as two major contributors to the global crisis of declining amphibian populations. These two factors have also been linked in explanations of the causes of these population declines. We conducted a combined exposure experiment to test the hypothesis that exposure to two agricultural herbicides would increase the susceptibility of post-metamorphic northern leopard frogs (Lithobates pipiens) to the amphibian fungal pathogen Batrachochytrium dendrobatidis (Bd). We assessed the independent and interactive effects of these exposures on the health and survival of the frogs. Wild-caught frogs underwent a 21-day exposure to a nominal concentration of either 2.1 µg/L atrazine (Aatrex(®) Liquid 480) or 100 µg a.e./L glyphosate (Roundup(®) Original), followed by Bd, and then were observed until 94 days post-initial exposure to the herbicides. Actual levels of atrazine were between 4.28 ± 0.04 µg/L and 1.70 ± 0.26 µg/L while glyphosate degraded from 100 µg a.e./L to approximately 7 µg a.e./L within 6 days of initial exposure to the herbicides. Compared to controls, the glyphosate formulation reduced the snout-vent length of frogs during the pesticide exposure (at Day 21), and the atrazine formulation reduced gain in mass up to Day 94. No treatment affected survival, splenosomatic or hepatosomatic indices, the densities and sizes of hepatic and splenic melanomacrophage aggregates, the density and size of hepatic granulomas, proportions of circulating leucocytes, the ratio of neutrophils to lymphocytes, or the ratio of leucocytes to erythrocytes. Histological assessment of samples collected at Day 94 revealed no evidence of Bd infection in any Bd-exposed frogs, while real-time PCR detected only one case of light infection in a single atrazine- and Bd-exposed frog. Frogs exposed to Bd shed their skin significantly more frequently than Bd-unexposed frogs, which may have helped them resist or clear infection, and could explain why no interaction between the herbicides and Bd was detected. The results suggest that these frogs were resistant to Bd infection and that pre-exposure to the herbicides did not alter this resistance. The effects seen on the growth following herbicide exposure is a concern, as reduced growth can lower the reproductive success and survival of the amphibians.

RECRUITMENT AND SEASONAL OCCURRENCE OF PARASITES IN JUVENILE INVASIVE ROUND GOBIES (NEOGOBIUS MELANOSTOMUS) IN THE ST. LAWRENCE RIVER, QUEBEC, CANADA.

The round goby (Neogobius melanostomus) is an invasive species that has become one of the most abundant fish in the St. Lawrence River, Quebec, Canada over the past 15 yr. Since its introduction, the round goby has acquired a number of native parasites, yet little is known about the dynamics of parasite recruitment. To examine this question, young-of-the-year and juvenile round gobies were collected monthly from 2 localities in the river (Îles de la Paix, Île Dorval) from June through November 2012. At Îles de la Paix, round gobies (n = 180) were infected with 3 species of parasites, all larval stages (Diplostomum spp., Tylodelphys scheuringi, Neoechinorhynchus tenellus). Prevalence of the digenean Diplostomum spp. varied from 3.3 to 13.3%, and mean abundance from 0.03 to 0.53 from June through September, with a maximum in August. The digenean T. scheuringi was seen only in August, at a prevalence of 10.0% and a mean abundance of 0.53. The acanthocephalan N. tenellus was observed in June, August, and September, prevalence ranging from 3.3 to 10.0% and mean abundance from 0.03 to 0.27. Maximum infection for all 3 species occurred in August. All infected fish were ≥44 mm in total length (TL). Fish infected with more than 1 parasite species were >60 mm TL. No round goby (n = 178) was infected at Île Dorval. This study demonstrated that the invasive round goby starts to acquire parasite infections in the St. Lawrence River in the first year of life and may contribute to the transmission of some parasites within this ecosystem.

Estimating the feeding range of a mobile consumer in a river-flood plain system using δ(13)C gradients and parasites.

1. The feeding range of an individual is central to food web dynamics as it determines the spatial scale of predator-prey interactions. However, despite recognition of its importance as a driving force in population dynamics, establishing feeding range is seldom done as detailed information on trophic interactions is difficult to obtain. 2. Biological markers are useful to answer this challenge as long as spatial heterogeneity in signal is present within the area investigated. A spatially complex ecosystem, Lake St. Pierre (LSP), a fluvial lake of the St Lawrence River (Québec, Canada), offered a unique opportunity to determine the feeding range of a secondary consumer, yellow perch (Perca flavescens) using isotopic ratios of carbon (δ(13)C). However, because food chains based on phytoplankton have generally more negative δ(13) C than those depending on periphyton, it was essential to determine the contribution of zooplankton in fish diet to correctly interpret spatial patterns of δ(13)C. We used parasites in perch to examine whether their δ(13)C was reflecting local δ(13)C baseline conditions rather than a feeding specialization on zooplankton. 3. δ(13)C of primary consumers was highly variable and exhibited a striking gradient along the shore-channel axis, suggesting that δ(13)C should reflect an individual consumer's spatial position in LSP. 4. This strong isotopic gradient allowed us to estimate the spatial scale of the resources used by individual perch following an approach presented by Rasmussen, Trudeau & Morinville (Journal of Animal Ecology, 78, 2009, 674). By comparing the δ(13)C variability in perch to that of primary consumers, we estimated that the adults feeding range was around 2 km along the shore-channel axis. 5. The combined use of isotopic ratios and parasites allowed us to determine that the adult population uses a wide range of habitats between the flood plain and the main channel. However, individually, each perch depended on a limited foodshed.

Natural and anthropogenic factors shape metazoan parasite community structure in mummichog (Fundulus heteroclitus) from two estuaries in New Brunswick, Canada.

Previously published multidisciplinary studies in the Miramichi and Bouctouche rivers (New Brunswick, Canada) noted significant changes in fish health parameters, including elevated tissue levels of organic contaminants and a wide range of physiological disturbances, in mummichog Fundulus heteroclitus (L.) from a site on the Miramichi River that received bleached kraft pulpmill and municipal effluent. The present study reports differences in the abundance of individual parasite species, as well as parasite infracommunity and component community composition, in mummichog from both rivers. These differences were evaluated in relation to host (size, condition, immune function, tissue organochlorine contaminant levels) and environmental (faecal coliform counts, salinity, temperature) data derived from the previously published studies. Overall, 18 parasite species were identified, the most common of which were Ascocotyle sp. larv., Ornithodiplostomum sp. larv., Posthodiplostomum sp. larv., and Proteocephalus filicollis (Rudolphi, 1802). There were broad differences in parasite community structure and composition between rivers and within rivers, the most prominent pattern being a pronounced difference between sites in the upper and lower estuary of each river that was likely driven by salinity. Mean infracommunity richness was also positively related to faecal coliforms (considered here as a surrogate measure of eutrophication via municipal sewage), and both were highest at the most polluted site. We noted no other significant relationships. Thus our data suggest that the parasite communities in these two estuaries were primarily structured by large upstream/downstream ecological gradients in salinity, and secondarily by eutrophication due to pollution by municipal and industrial effluents. Overall, our results highlight the value of coordinated multidisciplinary studies for understanding the factors that shape parasite abundance and community structure.

Myxozoan Communities in Two Cyprinid Fishes from Mesotrophic and Eutrophic Rivers.

Few studies have examined community structure among myxozoan species in fish. Herein myxozoan communities are described from 2 cyprinid species, the spottail shiner (Notropis hudsonius) and the common shiner (Luxilus cornutus), from mesotrophic and eutrophic localities in rivers in southwestern Quebec, Canada. Four myxozoan species were found, and total prevalence ranged from 60 to 86.7% in spottail shiners at 4 localities along the Richelieu River. Component species richness ranged from 2 to 4 and mean infracommunity richness from 0.87 to 1.47. Prevalence, component species richness, and infracommunity species richness in the spottail shiner were comparable to those from other localities in the Great Lakes and St. Lawrence River that were exposed to municipal effluents. Nine myxozoan species were found in common shiners from the Bras d'Henri micro-watershed. Component species richness varied from 6 to 8 at 4 localities, with total prevalence being 100% at all localities except 1, where it was 80%. Mean infracommunity richness ranged from 1.73 to 2.27. Prevalence, component species richness, and infracommunity species richness in the common shiner from the Bras d'Henri micro-watershed were among the highest observed for myxozoan communities from any host species to date. It is concluded that moderate levels of eutrophication are sufficient to generate species-rich communities of myxozoan parasites in fishes.

DNA barcodes show cryptic diversity and a potential physiological basis for host specificity among Diplostomoidea (Platyhelminthes: Digenea) parasitizing freshwater fishes in the St. Lawrence River, Canada.

Diplostomoid metacercariae parasitize freshwater fishes worldwide and cannot be identified to species based on morphology. In this study, sequences of the barcode region of cytochrome c oxidase subunit 1 (CO1) were used to discriminate species in 1088 diplostomoids, most of which were metacercariae from fish collected in the St. Lawrence River, Canada. Forty-seven diplostomoid species were detected, representing a large increase in known diversity. Most species suggested by CO1 sequences were supported by sequences of internal transcribed spacer (ITS) of rDNA and host and tissue specificity. Three lines of evidence indicate that physiological incompatibility between host and parasite is a more important determinant of host specificity than ecological separation of hosts and parasites in this important group of freshwater fish pathogens. First, nearly all diplostomoid species residing outside the lens of the eyes of fish are highly host specific, while all species that occur inside the lens are generalists. This can be plausibly explained by a physiological mechanism, namely the lack of an effective immune response in the lens. Second, the distribution of diplostomoid species among fish taxa reflected the phylogenetic relationships of host species rather than their ecological similarities. Third, the same patterns of host specificity were observed in separate, ecologically distinctive fish communities.

Vertebrate diets derived from trophically transmitted fish parasites in the Bothnian Bay.

Parasites that are transmitted through predator-prey interactions may be used as indicators of trophic relationships between organisms. Yet, they are rarely used as such in the construction of topological (predator-prey) food webs. We constructed food webs of vertebrate trophic interactions using observed diet alone, trophically transmitted parasites alone, and the combination of the two based on data from 31 species of fish from the Bothnian Bay, Finland. The fish food web contained 530 links derived from observed diet, 724 links inferred from parasitism, and 1,058 links calculated from a combination of both stomach contents and parasites. This sub-web constructed from stomach contents had a mean of 17.1 links per fish species, while that using parasites had 23.4 links per fish. Combining the two diet indicators yielded 34.1 links per fish species, illustrating the complementarity of the two methods. Mean number of prey species per fish species was 12.5 using observed diet items, 15.8 using parasites, and 24.5 using both measures together. Mean number of predators per fish species was 7.4 using observed diet, 11.7 using parasites and 15.0 using both. A positive correlation was found between the mean number of parasites and the number of prey taxa in the diet among the fishes. Omnivorous fish had the highest diversity of both parasite species and prey items, while benthophagous fish had among the lowest. Mean total abundance and mean total prevalence of parasites correlated positively with fish size, with piscivores being the largest with the highest abundance and prevalence, while planktivores and benthivores had the lowest. Trophically transmitted parasites may be used to help construct vertebrate sub-webs and derive information about food web processes. Parasites alone provided equivalent if not more information than observed diet. However, resolution is improved by using parasites and observed diet together.

Parasite fauna of Etheostoma nigrum (Percidae: Etheostomatinae) in localities of varying pollution stress in the St. Lawrence River, Quebec, Canada.

Parasite communities were examined in johnny darters (Etheostoma nigrum) collected from five localities in the St. Lawrence River in southwestern Quebec: two reference localities, one polluted locality upstream of the Island of Montreal and downstream of industrial and agricultural activity, and two polluted localities downstream of the Island of Montreal in the plume from the wastewater treatment facility. Twenty-four helminth species were found. Fish from the upstream polluted locality had the highest parasite species richness and total parasite numbers, and fish from the downstream polluted localities the lowest. Nonmetric multivariate analyses were conducted using square-root-transformed Bray-Curtis dissimilarity index. An analysis of similarity, dendrogram of centroids, and a permutational multivariate analysis of variance with contrasts all showed that fish from the reference localities had different parasite community composition than those from the polluted localities, and fish from the upstream polluted locality had different parasite communities than fish from the downstream polluted localities. Differences between reference and polluted localities were mainly due to higher abundances of the brain-encysting trematode, Ornithodiplostomum sp., at the reference localities. Differences between upstream and downstream polluted localities were mainly due to a higher diversity and abundance of trematodes in fish at the upstream locality.

Why do parasitized hosts look different? Resolving the "chicken-egg" dilemma.

Phenotypic differences between infected and non-infected hosts are often assumed to be the consequence of parasite infection. However, pre-existing differences in hosts' phenotypes may promote differential susceptibility to infection. The phenotypic variability observed within the host population may therefore be a cause rather than a consequence of infection. In this study, we aimed at disentangling the causes and the consequences of parasite infection by calculating the value of a phenotypic trait (i.e., the growth rate) of the hosts both before and after infection occurred. That procedure was applied to two natural systems of host-parasite interactions. In the first system, the infection level of an ectoparasite (Tracheliastes polycolpus) decreases the growth rate of its fish host (the rostrum dace, Leuciscus leuciscus). Reciprocally, this same phenotypic trait before infection modulated the future level of host sensitivity to the direct pathogenic effect of the parasite, namely the level of fin degradation. In the second model, causes and consequences linked the growth rate of the fish host (the rainbow smelt, Osmerus mordax) and the level of endoparasite infection (Proteocephalus tetrastomus). Indeed, the host's growth rate before infection determined the number of parasites later in life, and the parasite biovolume then decreased the host's growth rate of heavily infected hosts. We demonstrated that reciprocal effects between host phenotypes and parasite infection can occur simultaneously in the wild, and that the observed variation in the host phenotype population was not necessarily a consequence of parasite infection. Disentangling the causality of host-parasite interactions should contribute substantially to evaluating the role of parasites in ecological and evolutionary processes.