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In social vespine wasps, the nest characteristics and associates vary intra- and interspecifically. I studied the nest characteristics and nest associates of three Dolichovespula wasp species in boreal Finland. The average number of combs and cells per nest was similar among D. media, D. saxonica and D. norwegica, and similar to Dolichovespula in the temperate zone. Most D. media and D. saxonica colonies produced both gynes and males, whereas D. norwegica produced mainly either one. The average colony lifespan of D. saxonica was a bit over two months, i.e. shorter than documented for Dolichovespula in the temperate zone. The proportion of nests parasitised by the ichneumonid wasp Sphecophaga vesparum was 36% in D. saxonica and 50% in D. media; S. vesparum has not been recorded from D. media before. The proportion of parasitized cells was 0.6–11% for D. saxonica and 0.2–22% for D. media. The proportion yellow, overwintering cocoons of S. vesparum was highly variable among the nests. The proportion of nests parasitised by the moth Aphomia sociella was 36% in D. saxonica and 13% in D. media. Sphecophaga and Aphomia were not recorded from D. norwegica colonies, which supports studies in the temperate zone. The study adds up to the previous research on the nest structure and nest associates of vespines, and together they indicate that there is geographical variation within Europe and globally in the host use of Sphecophaga.

Abstract— Variability in infection rates of trematodes Diplostomum pseudospathaceum in 0+ rainbow trout Oncorhynchus mykiss under different hydraulic regimes was studied in the experimental setup with regulated flow rates and environmental heterogeneity. The average infection rate in the conditions of the current was 1.5 times less than that in the stagnant water. An increase of the flow rate from 3.2 to 11.3 cm/s did not result in lower infection rates, while the interindividual variability in the infection rate (coefficient of variation) tended to increase along with changing the still-water conditions (20%) to the high flow rate environments (40%) within the experiment. A decrease in the average infection rate and an increase in the variability within the heterogenous flow of water could indicate the fact that some fish effectively avoid infection. It may be caused by redistribution of cercariae suspended in the water column across microhabitats with different flow rates together with fish behavior which allows them to choose sites with low parasite concentration.

Global warming is likely to lengthen the seasonal duration of larval release by parasites. We exposed freshwater mussel hosts, Anodonta anatina, from 2 high-latitude populations to high, intermediate and low temperatures throughout the annual cercarial shedding period of the sympatric trematodes Rhipidocotyle fennica and R. campanula, sharing the same transmission pathway. At the individual host level, under warmer conditions, the timing of the cercarial release in both parasite species shifted towards seasonally earlier period while its duration did not change. At the host population level, evidence for the lengthening of larvae shedding period with warming was found for R. fennica. R. campanula started the cercarial release seasonally clearly earlier, and at a lower temperature, than R. fennica. Furthermore, the proportion of mussels shedding cercariae increased, while day-degrees required to start the cercariae shedding decreased in high-temperature treatment in R. fennica. In R. campanula these effects were not found, suggesting that warming can benefit more R. fennica. These results do not completely support the view that climate warming would invariably increase the seasonal duration of larval shedding by parasites, but emphasizes species-specific differences in temperature-dependence and in seasonality of cercarial release.

Bipartite network analysis is a powerful tool to study the processes structuring interactions in ecological communities. In applying the method, it is assumed that the sampled interactions provide an accurate representation of the actual community. However, acquiring a representative sample may be difficult as not all species are equally abundant or easily identifiable. Two potential sampling issues can compromise the conclusions of bipartite network analyses: failure to capture the full range of interactions (sampling completeness) and use of a taxonomic level higher than species to evaluate the network (taxonomic resolution). We asked how commonly used descriptors of bipartite antagonistic communities (modularity, nestedness, connectance, and specialization [H2′]) are affected by reduced host sampling completeness, parasite taxonomic resolution, and their crossed effect, as they are likely to co-occur. We used a quantitative niche model to generate weighted bipartite networks that resembled natural host–parasite communities. The descriptors were more sensitive to uncertainty in parasite taxonomic resolution than to host sampling completeness. When only 10% of parasite taxonomic resolution was retained, modularity and specialization decreased by ~76% and ~12%, respectively, and nestedness and connectance increased by ~114% and ~345% respectively. The loss of taxonomic resolution led to a wide range of possible communities, which made it difficult to predict its effects on a given network. With regards to host sampling completeness, standardized nestedness, connectance, and specialization were robust, whereas modularity was sensitive (~30% decrease). The combination of both sampling issues had an additive effect on modularity. In communities with low effort for both sampling issues (50%–10% of sampling completeness and taxonomic resolution), estimators of modularity, and nestedness could not be distinguished from those of random assemblages. Thus, the categorical description of communities with low sampling effort (e.g., if a community is modular or not) should be done with caution. We recommend evaluating both sampling completeness and taxonomic certainty when conducting bipartite network analyses. Care should also be exercised when using nonrobust descriptors (the four descriptors for parasite taxonomic resolution; modularity for host sampling completeness) when sampling issues are likely to affect a dataset., Ecology, 104 (4), ISSN:0012-9658, ISSN:1939-9170

Interactions among symbiotic organisms and their hosts are major drivers of ecological and evolutionary processes. Monitoring the infection patterns among natural populations and identifying factors affecting these interactions are critical for understanding symbiont–host relationships. However, many of these interactions remain understudied since the knowledge about the symbiont species is lacking, which hinders the development of appropriate tools. In this study, we developed a digital droplet PCR (ddPCR) assay based on apicomplexan COX1 gene to detect an undescribed agamococcidian symbiont. We show that the method gives precise and reproducible results and enables detecting cryptic symbionts in low target concentration. We further exemplify the assay's use to survey seasonally sampled natural host (Pygospio elegans) populations for symbiont infection dynamics. We found that symbiont prevalence differs spatially but does not show seasonal changes. Infection load differed between populations and was low in spring and significantly increased towards fall in all populations. We also found that the symbiont prevalence is affected by host length and population density. Larger hosts were more likely to be infected, and high host densities were found to have a lower probability of infection. The observed variations could be due to characteristics of both symbiont and host biology, especially the seasonal variation in encounter rates. Our findings show that the developed ddPCR assay is a robust tool for detecting undescribed symbionts that are otherwise difficult to quantify, enabling further insight into the impact cryptic symbionts have on their hosts., Many symbiont‐host interactions are understudied due to the lack of knowledge about the symbiont species hindering the development of appropriate tools. Here, we developed a ddPCR assay to detect an undescribed agamococcidian symbiont in its host, Pygospio elegans, and show that the method enables detecting cryptic symbionts even in low target concentration. We further exemplify the assay's use to survey seasonally sampled natural host populations for symbiont infection dynamics.

Genetic variation in resistance against parasite infections is a predominant feature in host–parasite systems. However, mechanisms maintaining genetic polymorphism in resistance in natural host populations are generally poorly known. We explored whether differences in natural infection pressure between resource‐based morphs of Arctic charr (Salvelinus alpinus) have resulted in differentiation in resistance profiles. We experimentally exposed offspring of two morphs from Lake Þingvallavatn (Iceland), the pelagic planktivorous charr (“murta”) and the large benthivorous charr (“kuðungableikja”), to their common parasite, eye fluke Diplostomum baeri, infecting the eye humor. We found that there were no differences in resistance between the morphs, but clear differences among families within each morph. Moreover, we found suggestive evidence of resistance of offspring within families being positively correlated with the parasite load of the father, but not with that of the mother. Our results suggest that the inherited basis of parasite resistance in this system is likely to be related to variation among host individuals within each morph rather than ecological factors driving divergent resistance profiles at morph level. Overall, this may have implications for evolution of resistance through processes such as sexual selection., Mechanisms maintaining genetic polymorphism in resistance in natural host populations are generally poorly known. Here, we show no differences in resistance between sympatric morphs of Arctic charr, but clear differences among families within each morph. Our results suggest that the inherited basis of parasite resistance in this system is related to variation among host individuals within each morph rather than ecological factors driving divergent resistance profiles at morph level.

Temperature and intraspecific competition are important factors influencing the growth of all organisms, including parasites. The temperature increase is suggested to stimulate the development of parasites within poikilothermic hosts. However, at high parasite densities, this effect could be diminished, due to stronger intraspecific competition. Our study, for the first time, addressed the joint effects of warming and parasite abundances on parasite growth in poikilothermic hosts. The growth of the common fish parasite larvae (trematode Diplostomum pseudospathaceum) within the rainbow trout at different infection intensities and temperatures (15°C and 18°C) was experimentally investigated. The results showed that temperature was positively correlated with both parasite infection success and growth rates. The growth rates increased much more compared to those in many free-living poikilothermic animals. Atypically for a majority of parasites, D. pseudospathaceum larvae grow faster when abundant (Allee effect). The possible causes for this phenomenon (manipulation cost sharing, etc.) are discussed in this study. Importantly, limited evidence of the interaction between temperature and population density was found. It is likely that temperature did not change the magnitude of the Allee effect but affected its timing. The impact of these effects is supposed to become more pronounced in freshwater ecosystems under current climate changes.

Parasitoids and predators can cause marked mortality in their host species. I studied the occurrence, abundance, and biology of the wasp nest beetle Metoecus paradoxus (Ripiphoridae), inhabiting the nests of the common wasp Vespula vulgaris (Vespidae) in Central Finland. I also compiled phenological data of the species in Finland. The proportion of parasitized nests was 80%. The abundance of the beetle was generally some tens of individuals (max. = 130) per nest, and the parasitism rate 1–18% of the sealed cells, which are typical for the species in temperate Europe. The beetle occurred in several combs but was absent or rare in the oldest and newest combs. Sex ratio was female biased. Females were larger than males, and individuals from large cells were larger than from small cells. Adult beetles were observed rather steadily from early August to late September. The study shows that M. paradoxus is a common inhabitant of V. vulgaris nests in Central Finland, but due to a low frequency of parasitized cells its impact on wasp populations is likely to be minor.

Three subspecies of the ringed seal (Pusa hispida) are found in northeastern Europe: P. h. botnica in the Baltic Sea, P. h saimensis in Lake Saimaa in Finland, and P. h. ladogensis in Lake Ladoga in Russia. We investigated the poorly-known cestode helminth communities of these closely related but ecologically divergent subspecies using COI barcode data. Our results show that, while cestodes from the Baltic Sea represent Schistocephalus solidus, all worms from the two lakes are identified as Ligula intestinalis, a species that has previously not been reported from seals. The observed shift in cestode communities appears to be driven by differential availability of intermediate fish host species in marine vs. freshwater environments. Both observed cestode species normally infect fish-eating birds, so further work is required to elucidate the health and conservation implications of cestode infections in European ringed seals, whether L. intestinalis occurs also in marine ringed seals, and whether the species is able to reproduce in seal hosts. In addition, a deep barcode divergence found within S. solidus suggests the presence of cryptic diversity under this species name., Graphical abstract Image 1, Highlights • COI barcoding reveals different cestodes in marine and freshwater ringed seals. • Ligula intestinalis is reported for the first time from seals. • A deep barcode divergence is found within Schistocephalus solidus in the Baltic Sea.

Infectious diseases are key drivers of wildlife populations and agriculture production, but whether and how climate change will influence disease impacts remains controversial. One of the critical knowledge gaps that prevents resolution of this controversy is a lack of high-quality experimental data, especially in marine systems of significant ecological and economic consequence. Here, we performed a manipulative experiment in which we tested the temperature-dependent effects on Atlantic salmon (Salmo salar) of sea lice (Lepeophtheirus salmonis)—a parasite that can depress the productivity of wild-salmon populations and the profits of the salmon-farming industry. We explored sea-louse impacts on their hosts across a range of temperatures (10, 13, 16, 19, and 22 °C) and infestation levels (zero, ‘low’ (mean abundance ± SE = 1.6 ± 0.1 lice per fish), and ‘high’ infestation (6.8 ± 0.4 lice per fish)). We found that the effects of sea lice on the growth rate, condition, and survival of juvenile Atlantic salmon all worsen with increasing temperature. Our results provide a rare empirical example of how climate change may influence the impacts of marine disease in a key social-ecological system. These findings underscore the importance of considering climate-driven changes to disease impacts in wildlife conservation and agriculture.

A moderate raise in temperature was suggested to enhance the impact of parasites on aquatic ecosystems. Under higher temperatures, poikilothermic animals (e.g. fish) increase their activity, which can result in a more frequent encounter with parasites. However, temperature increase may also trigger processes counteracting an increased risk of parasitic infections. Thus, the removal of free‐living stages of parasites by filter‐feeding organisms can increase with temperature and potentially mitigate disease risk in ecosystems under climate change. We aimed to study whether an increased infection transmission under higher temperatures can be compensated by the increased removal of parasitic larvae by aquatic predators. In addition, we planned to reveal the behavioral mechanism underlying the more successful transmission of the parasite at higher temperatures. We studied experimentally how temperature, the behavior of fish hosts (rainbow trout), and the presence of filter‐feeding mussels in the environment influence the infection success of trematode larvae Diplostomum pseudospathaceum cercariae. We found that temperature raise increased while the presence of filter‐feeding mussels in the environment decreased infection intensities in fish. However, the effect of mussel's presence was constant within the tested range of water temperatures (15–23°C), which suggests that it cannot compensate for the observed increased transmission of parasites under temperature raise. Fish activity before the exposure to parasites was a substantial factor affecting the host's vulnerability to infection. However, fish activity only weakly correlated with temperature, therefore, it is unlikely to be the only factor responsible for the increased infection success under warmer conditions. After the exposure, fish activity decreased and did not influence the infection's success. This decrease was temperature‐dependent and more pronounced in more active fish. In general, we showed that the elimination of trematode larvae by filter‐feeders is unlikely to deter the potential effects of global warming on host–parasite interactions in temperate freshwater ecosystems. peerReviewed

Coinfections of two or more parasites within one host are more of a rule than an exception in nature. Interactions between coinfecting parasites can greatly affect their abundance and prevalence. Characteristics of the host, such as genetic diversity, can also affect the infection dynamics of coinfecting parasites. Here we investigate for the first time the association of coinfection patterns of two marine apicomplexans, Rhytidocystis sp. and Selenidium pygospionis, with the genetic diversity of their host, the polychaete Pygospio elegans, from natural populations. Host genetic diversity was determined with seven microsatellite loci and summarized as allelic richness, inbreeding coefficient, and individual heterozygosity. We detected nonsignificant correlations between infection loads and both individual host heterozygosity and population genetic diversity. Prevalence and infection load of Rhytidocystis sp. were higher than that of S. pygospionis and both varied spatially. Coinfections were common and almost all hosts infected by S. pygospionis were also infected by Rhytidocystis sp.. Rhytidocystis sp. infection load was significantly higher in dual infections. Our results suggest that factors other than host genetic diversity might be more important in marine apicomplexan infection patterns and experimental approaches would be needed to further determine how interactions between the apicomplexans and their host influence infection. peerReviewed

Several northern freshwater fishes have gone through rapid adaptive radiation after the last glacial period, resulting in new species or intraspecific morphs with distinct life histories. Parasite infections can promote adaptive radiations and spatiotemporal differences in patterns of infections can potentially reveal incipient or ongoing speciation processes. We investigated intraspecific differentiation in a freshwater gadoid fish, burbot (Lota lota), by exploring differences in parasite infections between two potential life-history morphs in Lake Konnevesi, Central-Finland, one reproducing species characteristically in shallow littoral waters in February and the other possibly in deep profundal zone roughly a month later. By conducting a sampling campaign on reproducing fish over two consecutive years, we found significant differences in infections between the fish captured from littoral and profundal sites. More specifically, larval trematode and cestode infections were consistently less abundant in profundal fish, tracking long-term exposure differences in shallow waters. In contrast, trophically transmitted metazoan infections in the fish gut, reflecting shorter-term differences in feeding, showed higher variation between sampling years rather than depths. We also found suggestive evidence of higher trematode-inflicted tissue damage per parasite in the profundal fish, implying lower tolerance to the infection. These results offer further evidence that burbot captured from littoral and profundal sites represent differentiated life-history morphs. We propose that ecological and evolutionary differentiation within burbot populations across its circumpolar distribution may be more widespread than previously acknowledged. peerReviewed

Parasitic salmon lice (Lepeophtheirus salmonis) threaten the economic and ecological sustainability of salmon farming, and their evolved resistance to treatment with emamectin benzoate (EMB) has been a major problem for salmon farming in the Atlantic Ocean. In contrast, the Pacific Ocean, where wild salmon are far more abundant, has not seen widespread evolution of EMB-resistant lice. Here, we use EMB bioassays and counts of lice on farms from the Broughton Archipelago, Canada—a core region of salmon farming in the Pacific—to show that EMB sensitivity has dramatically decreased since 2010, concurrent with marked decrease in the field efficacy of EMB treatments. Notably, these bioassay data were not made available through public reporting by industry or by the federal regulator, but rather through Indigenous-led agreements that created a legal obligation for salmon-farming companies to provide data to First Nations. Our results suggest that salmon lice in the Pacific Ocean have recently evolved substantial resistance to EMB, and that salmon-louse outbreaks on Pacific farms will therefore be more difficult to control in the coming years.

Effect of freshwater mussels’ (Unionoida) glochidia on the growth of fish host has remained poorly studied. We compared the specific growth rate of the juvenile, PIT-marked brown trout (Salmo trutta) between uninfected controls to those experimentally infected (average initial intensity of infection 8000 fish−1) with Margaritifera margaritifera glochidia, kept in high and low feeding. Growth and mortality of fish were monitored for 10 months. Our hypothesis was that glochidiosis would impair the growth of fish. According to our hypothesis, infected fish gained statistically significantly less weight than the control fish throughout the experiment. A proportional increase in weight of control individuals was 11% higher than that of the infected fish. However, neither the feeding regime (high, low) nor the period (September–November, November–March, March–May), had a significant effect on the growth difference between control and infected fish. As the effect of infection on the growth of fish was subtle and no effect on host mortality was detected either, this may turn public opinion favorable for M. margaritifera conservation even if the salmonid host population is important for commercial or recreational fishing.

The reduction in host fitness caused by parasite infections (virulence) depends on infection intensity and the degree of damage caused per parasite. Environmental conditions can shape both virulence components, but in contrast to infection intensity, environmental impacts on per-parasite damage are poorly understood. Here, we studied the effect of ambient temperature on per-parasite damage, which is jointly determined by the ability of parasites to induce harm (per-parasite pathogenicity) and the ability of hosts to limit damage (tolerance). We experimentally exposed two salmonid species, Atlantic salmon (Salmo salar) and sea trout (Salmo trutta), to replicated genotypes of the eye fluke Diplostomum pseudospathaceum. After development of health damage (eye cataracts) in warm water (16 °C) during the first 12 weeks post exposure, we maintained the fish at either 5 °C (cold water) or 16 °C for another 8 weeks and quantified changes in cataracts as a function of parasite load. We found that per-parasite damage was reduced in cold compared to warm water, suggesting that cold temperatures improved host health. Per-parasite damage was also affected by parasite genotype and host species, but these effects did not change with temperature. Our findings suggest that cold-water seasons, which are often neglected in host-parasite studies due to low infection risk, could allow hosts to recuperate and thus, may have important implications for the ecology and epidemiology of parasite infections. peerReviewed

As a consequence of climate change and open net-pen salmon farming, wild Atlantic salmon Salmo salar are increasingly likely to encounter elevated temperatures and parasite abundances during their early marine migration. Such stressors can compromise fitness by diminishing liver energy stores and impairing cardiac muscle. To assess whether temperature and infestation by salmon lice Lepeophtheirus salmonis are important correlates of liver energy stores and cardiac muscle performance in juvenile salmon, we experimentally infested fish at 3 abundances of louse infestation (zero, low, and high) and 5 temperatures (10, 13, 16, 19, and 22°C). At the end of the experiment (i.e. when sea lice reached adulthood), we calculated the percent dry weight of the liver (%DWL; a proxy for liver energy stores) and cardiosomatic index (CSI; a proxy for cardiac muscle performance) of each fish and fitted 5 linear mixed-effects models to both of these responses. For both %DWL and CSI, the best-supported model included additive fixed effects for both infestation level and temperature. Our top models predicted that, relative to zero infestation, high infestation reduces %DWL by 5.7% (95% CI: 5.3-6.2%) and increases CSI by 15.9% (14.4-18.0%), and low infestation reduces %DWL by 2.6% (2.2-3.0%) and increases CSI by 7.8% (6.7-10.0%). Our work suggests that stressors associated with ocean warming and coastal salmon aquaculture can compromise wild salmon fitness through the impairment of vital organs. peerReviewed

An increasing threat to local, native freshwater mussels (Unionida)—an ecologically important but globally alarmingly declining group— is the invasion by exotic bivalves. The Enemy Release Hypothesis predicts that introduced species should benefit from enemy-mediated competition because they are less likely to be harmed by natural enemies, such as parasites, than their native competitors. We investigated within-site differences in parasitism between sympatric native (tot. five spp.) and invasive (tot. three spp.) bivalves in eight northern European waterbodies, which harboured totally 15 parasite taxa. In paired comparisons using within-site averages, the mean number of parasite species in the native bivalves was 2.3 times higher, and the sum of parasite prevalences 2.4 times higher, than in the invasive bivalves. This may lead to enemy-mediated compet- itive release of invaders and contribute to the success of invasive freshwater bivalves, in general. However, while the invasive clam Corbicula fluminea was completely free from parasites, parasite parameters of the other invader, Sinanodonta woodiana, were relatively high, indicating that the role of parasites can be invader-specific and urges further research. Under- standing the factors affecting success of freshwater bivalve invasions, such as parasitism, can aid invasion control and conservation of local, native (endangered) bivalves. Open access funding provided by University of Jyväskylä (JYU). Co-authors’ cooperation was initiated with the support of the Poznan ́ City Council under the ‘‘Academic and Scientific Poznan ́’’ Program. The study was funded by Estonian Ministry of Education and Research (institutional research funding project IUT 21-2 to Tiina Nõges) and by Estonian Research Council, Mobilitas Pluss research project MOBJD29. This study was also supported by the Polish Minister of Science and Higher Education, under the program ‘‘Regional Initiative of Excellence’’ in 2019–2022 (Grant No. 008/RID/2018/19), by Poznan ́ University of Life Sciences, University of Jyva ̈skyla ̈, and by CNR Short Term Mobility Project 2016. We thank Henryk Gierszal and Henn Timm for valuable assistance in field and laboratory. Open access funding provided by University of Jyväskylä (JYU). Co-authors’ cooperation was initiated with the support of the Poznan ́ City Council under the ‘‘Academic and Scientific Poznan ́’’ Program. The study was funded by Estonian Ministry of Education and Research (institutional research funding project IUT 21-2 to Tiina Nõges) and by Estonian Research Council, Mobilitas Pluss research project MOBJD29. This study was also supported by the Polish Minister of Science and Higher Education, under the program ‘‘Regional Initiative of Excellence’’ in 2019–2022 (Grant No. 008/RID/2018/19), by Poznan ́ University of Life Sciences, University of Jyva ̈skyla ̈, and by CNR Short Term Mobility Project 2016. We thank Henryk Gierszal and Henn Timm for valuable assistance in field and laboratory.

Infections with microsporidian parasites are described in skeletal muscle of burbot Lota lota from Lake Haukivesi, Finland. Infected myocytes contained spores within sporophorous vesicles (SPVs) in contact with host cell cytoplasm, similar to Pleistophora ladogensis in L. lota and smelt Osmerus eperlanus in western Russia and northern Germany. Analysis of small subunit ribosomal RNA (SSU rRNA) gene sequences indicated identity with Myosporidium spraguei in burbot and pike-perch from this lake. The latter is considered a junior synonym of P. ladogensis. Phylogenetic analysis of SSU rRNA sequences resolved the burbot parasite apart from a clade containing the type species P. typicalis, but together with M. merluccius. The parasite is renamed Myosporidium ladogensis (Voronin, 1978) n. comb. Networks of tubular appendages arising from developing meronts and SPVs were associated with degradation of host cell cytoplasm.

There is increasing evidence that the composition of a social group influences the fitness of its members. For example, group member identities can determine the exposure risk to contact-transmitted parasites and consequently impact the health of all group members. Here, we propose that group composition may also affect host exposure to parasite propagules prevailing in the environment via collective parasite avoidance behaviours. We explored the spatial avoidance of a trematode parasite, Diplostomum pseudospathaceum, using the simplest form of host groups, pairs of sea trout, Salmo trutta trutta. These pairs showed either (1) between-group heterogeneity in their experience with the parasite (both, one or neither previously exposed) and matched dominance ranks (assessed through growth rates) or (2) within-group heterogeneity in previous exposure and in dominance ranks. We show that pairs that had recently been exposed to the parasite spent significantly less time in areas of infection risk than pairs in which one or both individuals were unexposed. This result suggests that previous exposure improves parasite avoidance. Among pairs with mixed exposure history, parasite avoidance was most pronounced when the exposed individual was dominant over its partner. We conclude that higher ranked individuals direct collective movements and their exposure history predicts a pair's parasite avoidance. As avoidance efficiency was directly related to infection rate, our results suggest that group composition can shape individual exposure to nonsocially transmitted parasites, and ultimately fitness.

Preinfection by one parasitic species may facilitate or by contrast hamper the subsequent penetration and/or establishment of other parasites in a host. The biology of interacting species, timing of preinfection, and dosage of subsequent parasite exposure are likely important variables in this multiparasite dynamic infection process. The increased vulnerability to subsequent infection can be an important and often overlooked factor influencing parasite virulence. We investigated how the preinfection by freshwater pearl mussel Margaritifera margaritifera glochidia could influence the success of subsequent infection by the common trematode Diplostomum pseudospathaceum in brown trout Salmo trutta and vice versa whether preinfection by the trematode made fish more susceptible to glochidia infection. The first experiment was repeated twice with different (low and high) exposure doses to initiate the subsequent trematode infection, while in the second experiment we varied the timing of the preinfection with trematodes. The preinfection with glochidia made fish more vulnerable to subsequent infection with trematodes. Since the trematodes penetrate through the gills, we suggest that increased host vulnerability was most likely the result of increased respiration caused by the freshwater pearl mussel glochidia encysted on gills. In turn, brown trout preinfected with trematodes were more vulnerable to the subsequent glochidial infection, but only if they were preinfected shortly before the subsequent infection (20 hr). Fish preinfected with trematodes earlier (2 weeks before the subsequent infection) did not differ in their vulnerability to glochidia. These effects were observed at moderate intensities of infections similar to those that occur in nature. Our study demonstrates how the timing and sequence of exposure to parasitic species can influence infection success in a host–multiparasite system. It indicates that the negative influence of glochidia on host fitness is likely to be underestimated and that this should be taken into consideration when organizing freshwater pearl mussel restoration procedures. peerReviewed

This book chapter describes various aspects of diplostomiasis caused by Diplostomum spathaceum: diagnosis, epidemiology, life cycle, transmission, developmental stages, population dynamics, effects of climate change on parasite distribution, and disease control and prevention. peerReviewed

Genetic variation in defence against parasite infections is fundamental for host–parasite evolution. The overall level of defence of a host individual or population includes mechanisms that reduce parasite exposure (avoidance), establishment (resistance) or pathogenicity (tolerance). However, how these traits operate and evolve in concert is not well understood. Here, we investigated genetic variation in and associations between avoidance, resistance and tolerance in a natural host–parasite system. Replicated populations of Atlantic salmon ( Salmo salar ) and sea trout (an anadromous form of brown trout, Salmo trutta ) were raised under common garden conditions and infected with the eye fluke Diplostomum pseudospathaceum . We demonstrate significant genetic variation in the defence traits across host populations and negative associations between the traits, with the most resistant populations showing the weakest avoidance and the lowest infection tolerance. These results are suggestive of trade-offs between different components of defence and possibly underlie the genetic variation in defence traits observed in the wild. Because the three defence mechanisms affect host–parasite evolution in profoundly different ways, we emphasize the importance of studying these traits in concert.

This is a post-peer-review, pre-copyedit version of an article published in Hydrobiologia. The final authenticated version is available online at https://doi.org/10.1007/s10750-019-3953-4. Niche diversification of polymorphic Arctic charr can be altered by multiple anthropogenic stressors. The opossum-shrimp (Mysis relicta) was introduced to compensate for reduced food resources for fish following hydropower operations in Lake Limingen, central Norway. Based on habitat use, stomach contents, stable isotopes (δ13C, δ15N) and trophically transmitted parasites, the zooplanktivorous upper water-column dwelling ‘normal’ morph was clearly trophically separated from two sympatric deep-water morphs (the ‘dwarf’ and the ‘grey’) that became more abundant with depth (> 30 m). Mysis dominated (50–60%) charr diets in deeper waters (> 30 m), irrespective of morph. Mysis and/or zooplankton prey groups caused high dietary overlap (> 54%) between the ‘dwarf’ morph and the two other ‘normal’ and ‘grey’ morphs. After excluding Mysis, the dietary overlap dropped to 34% between the two profundal morphs, as the ‘dwarf’ fed largely on deep-water zoobenthos (39%), while the ‘grey’ morph fed on fish (59%). The time-integrated trophic niche tracers (trophically transmitted parasites and stable isotopes) demonstrated only partial dietary segregation between the two deep-water morphs. The high importance of Mysis in Arctic charr diets may have reduced the ancestral niche segregation between the deep-water morphs and thereby increased their resource competition and potential risk of hybridization.

Summary Phytoplankton and bacteria interactions have a significant role in aquatic ecosystem functioning. Associations can range from mutualistic to parasitic, shaping biogeochemical cycles and having a direct influence on phytoplankton growth. How variations in phenotype and sampling location, affect the phytoplankton microbiome is largely unknown. A high‐resolution characterization of the bacterial community in cultures of the dinoflagellate Alexandrium was performed on strains isolated from different geographical locations and at varying anthropogenic impact levels. Microbiomes of Baltic Sea Alexandrium ostenfeldii isolates were dominated by Betaproteobacteria and were consistent over phenotypic and genotypic Alexandrium strain variation, resulting in identification of an A. ostenfeldii core microbiome. Comparisons with in situ bacterial communities showed that taxa found in this A. ostenfeldii core were specifically associated to dinoflagellate dynamics in the Baltic Sea. Microbiomes of Alexandrium tamarense and minutum, isolated from the Mediterranean Sea, differed from those of A. ostenfeldii in bacterial diversity and composition but displayed high consistency, and a core set of bacterial taxa was identified. This indicates that Alexandrium isolates with diverse phenotypes host predictable, species‐specific, core microbiomes reflecting the abiotic conditions from which they were isolated. These findings enable in‐depth studies of potential interactions occurring between Alexandrium and specific bacterial taxa.

Leakage of medical residues into the environment can significantly impact natural communities. For example, antibiotic contamination from agriculture and aquaculture can directly influence targeted pathogens, but also other non-targeted taxa of commensals and parasites that regularly co-occur and co-infect the same host. Consequently, antibiotics could significantly alter interspecific interactions and epidemiology of the co-infecting parasite community. We studied how minor environmental concentrations of antibiotic affects the co-infection of two parasites, the bacterium Flavobacterium columnare and the fluke Diplostomum pseudospathaceum , in their fish host. We found that antibiotic in feed, and particularly the minute concentration in water, significantly decreased bacterial virulence and changed the infection success of the flukes. These effects depended on the level of antibiotic resistance of the bacterial strains. Antibiotic, however, did not compensate for the higher virulence of co-infections. Our results demonstrate that even very low environmental concentrations of antibiotic can influence ecology and epidemiology of diseases in co-infection with non-targeted parasites. Leakage of antibiotics into the environment may thus have more complex effects on disease ecology than previously anticipated.

SUMMARYFactors that drive parasite specificity and differences in infection dynamics among alternative host species are important for ecology and evolution of host–parasite interactions, but still often poorly known in natural systems. Here, we investigated spatiotemporal dynamics of infection, host susceptibility and parasite-induced changes in host phenotype in a rarely explored host–parasite system, theAustralapatemonsp. trematode infecting two sympatric species of freshwater leeches,Erpobdella octoculataandHelobdella stagnalis. We show significant variation in infection abundance between the host species in both space and time. Using experimental infections, we also show that most of this variation likely comes from interspecific differences in exposure rather than susceptibility. Moreover, we demonstrate that the hiding behaviour ofE. octoculata, but not that ofH. stagnalis, was impaired by the infection irrespective of the parasite abundance. This may increase susceptibility ofE. octoculatato predation by the final avian host. We conclude that differences in patterns of infection and in behavioural alterations among alternative sympatric host species may arise in narrow spatial scales, which emphasises the importance of local infection and transmission dynamics for parasite life cycles.

Coinfections by multiple parasites predominate in the wild. Interactionsbetween parasites can be antagonistic, neutral, or facilitative, and they canhave significant implications for epidemiology, disease dynamics, and evolu-tion of virulence. Coinfections commonly result from sequential exposure ofhosts to different parasites. We argue that the sequential nature of coinfectionsis important for the consequences of infection in both natural and man-madeenvironments. Coinfections accumulate during host lifespan, determining thestructure of the parasite infracommunity. Interactions within the parasite com-munity and their joint effect on the host individual potentially shape evolution ofparasite life-history traits and transmission biology. Overall, sequential coin-fections have the potential to change evolutionary and epidemiological out-comes of host–parasite interactions widely across plant and animal systems. peerReviewed

Pelodera strongyloides is a generally free-living gonochoristic facultative nematode. The whole genomic sequence of P. strongyloides remains unknown but 4 small subunit ribosomal RNA (ssrRNA) gene sequences are available. This project launched a de novo transcriptome assembly with 100 bp paired-end RNA-seq reads from normal, starved and wet-plate cultured animals. Trinity assembly tool generated 104,634 transcript contigs with N50 contig being 2195 bp and average contig length at 1103 bp. Transcriptome BLASTX matching results of five nematodes (C. elegans, Strongyloides stercoralis, Necator americanus, Trichuris trichiura, and Pristionchus pacificus) were consistent with their evolutionary relationships. Sixteen genes were identified to be homologous to key elements of the C. elegans RNA interference system, such as Dicer, Argonaute, RNA-dependent RNA polymerase and double strand RNA transport proteins. In starved samples, we observed up-regulation of cuticle related genes and 3 dauer formation genes. Dauer morphology was captured with enlarged phasmid under light microscopy, and dauer and normal larvae counts in clumps had a Pearson's product-moment correlation of 0.805 with P-value = 0.0088. Our results demonstrate that P. strongyloides could be used for studying nematode-related human or pet parasitic diseases. The sequenced assembled transcriptome reported here may be useful to understand the evolution of parasitism in Nematoda. peerReviewed

Within‐host interactions between co‐infecting parasites can significantly influence the evolution of key parasite traits, such as virulence (pathogenicity of infection). The type of interaction is expected to predict the direction of selection, with antagonistic interactions favouring more virulent genotypes and synergistic interactions less virulent genotypes. Recently, it has been suggested that virulence can further be affected by the genetic identity of co‐infecting partners (G × G interactions), complicating predictions on disease dynamics. Here, we used a natural host–parasite system including a fish host and a trematode parasite to study the effects of G × G interactions on infection virulence. We exposed rainbow trout (Oncorhynchus mykiss) either to single genotypes or to mixtures of two genotypes of the eye fluke Diplostomum pseudospathaceum and estimated parasite infectivity (linearly related to pathogenicity of infection, measured as coverage of eye cataracts) and relative cataract coverage (controlled for infectivity). We found that both traits were associated with complex G × G interactions, including both increases and decreases from single infection to co‐infection, depending on the genotype combination. In particular, combinations where both genotypes had low average infectivity and relative cataract coverage in single infections benefited from co‐infection, while the pattern was opposite for genotypes with higher performance. Together, our results show that infection outcomes vary considerably between single and co‐infections and with the genetic identity of the co‐infecting parasites. This can result in variation in parasite fitness and consequently impact evolutionary dynamics of host–parasite interactions. peerReviewed

Parasites play a central role in the adaptiveness of sexual reproduction. Sexual selection theory suggests a role for parasite resistance in the context of mate choice, but the evidence is mixed. The parasite-mediated sexual selection (PMSS) hypothesis derives a number of predictions, among which that resistance to parasites is heritable, and that female choice favors parasite resistance genes in males. Here, we tested the PMSS hypothesis using the dung beetle Onthophagus taurus, a species that can be heavily parasitized by Macrocheles merdarius mites, which are known to affect adult survival. We investigated the heritability of resistance to M. merdarius, as well as whether female O. taurus impose a mating bias against males susceptible to mite infestation. Female choice for parasite resistance is difficult to disentangle from the possibility that females are simply choosing less parasitized males due to naturally selected benefits of avoiding contracting those parasites. This is especially likely for ectoparasites, such as mites. We tackled this problem by performing a mate choice trial first, and then measuring a male’s resistance to mite infestation. Resistance to mite infestation exhibited significant levels of additive genetic variance. Although we found no relationship between mating success and parasite resistance, males with greater resistance to infestation mated for longer. If females control copula duration, given that short copulations often result in mating failure, female choice could act on parasite resistance. peerReviewed

Influence of fish personality on infection rate is almost not studied. In the experiments on the young-of-the-year Oncorhynchus mykiss and cercariae of a trematode Diplostomum pseudospathaceum we tested the hypothesis that infection rate differs between more and less active (“bold” and “shy”) fish. Will individual differences in infection persist upon re-infection? Fish serve as a second intermediate host for this trematode. A positive correlation was found between the results of consecutive infections. Accumulation of parasites with successive infections leads to an aggregated distribution of D. pseudospathaceum among the hosts, affecting individual fitness and polymorphism in fish populations. Persistent individual differences in parasite burden among fish and, as a result, vulnerability for predators confirms the role of parasites as an important factor of natural selection. peerReviewed

Removal of parasite free-living stages by predators has previously been suggested an important factor controlling parasite transmission in aquatic habitats. Experimental studies of zooplankton predation on macroparasite larvae are, however, scarce. We tested whether trematode cercariae, which are often numerous in shallow waters, are suitable prey for syntopic zooplankters. Feeding rates and survival of freshwater cyclopoids (Megacyclops viridis, Macrocyclops distinctus), calanoids (Arctodiaptomus paulseni), cladocerans (Sida crystallina) and rotifers Asplanchna spp., fed with cercariae of Diplostomum pseudospathaceum, a common fish trematode, were studied. In additional long-term experiments, we studied reproduction of cyclopoids fed with cercariae. All tested zooplankton species consumed cercariae. The highest feeding rates were observed for cyclopoids (33 ± 12 cercariae ind−1 h−1), which actively reproduced (up to one egg clutch day−1) when fed ad libitum with cercariae. Their reproductive characteristics did not change significantly with time, indicating that cercariae supported cyclopoids’ dietary needs. Mortality of rotifers and cladocerans was high (25–28% individuals) when exposed to cercariae in contrast to cyclopoids and calanoids (

Studies on the determinants of plant-herbivore and herbivore-parasitoid associations provide important insights into the origin and maintenance of global and local species richness. If parasitoids are specialists on herbivore niches rather than on herbivore taxa, then alternating escape of herbivores into novel niches and delayed resource tracking by parasitoids could fuel diversification at both trophic levels. We used DNA barcoding to identify parasitoids that attack larvae of seven Pontania sawfly species that induce leaf galls on eight willow species growing in subarctic and arctic-alpine habitats in three geographic locations in northern Fennoscandia, and then applied distance- and model-based multivariate analyses and phylogenetic regression methods to evaluate the hierarchical importance of location, phylogeny and different galler niche dimensions on parasitoid host use. We found statistically significant variation in parasitoid communities across geographic locations and willow host species, but the differences were mainly quantitative due to extensive sharing of enemies among gallers within habitat types. By contrast, the divide between habitats defined two qualitatively different network compartments, because many common parasitoids exhibited strong habitat preference. Galler and parasitoid phylogenies did not explain associations, because distantly related arctic-alpine gallers were attacked by a species-poor enemy community dominated by two parasitoid species that most likely have independently tracked the gallers' evolutionary shifts into the novel habitat. Our results indicate that barcode- and phylogeny-based analyses of food webs that span forested vs. tundra or grassland environments could improve our understanding of vertical diversification effects in complex plant-herbivore-parasitoid networks.

Group‐living animals can gain protection against parasitic infections through social contacts with previously infected conspecifics (social immunization). Recent research suggests that such protective effects can be induced through visual or chemical cues released by infected individuals, resulting in anticipatory immune upregulation among group members. Here, we study cue‐induced social resistance in rainbow trout Oncorhynchus mykiss exposed to a trematode parasite, the eye‐fluke Diplostomum pseudospathaceum. We established groups of naïve individuals (receivers) that were paired with previously infected individuals (donors) at different ratios of donors to receivers and at different time points since donor exposure to capture varying concentrations of the anticipated cues. While the pre‐infection elevated resistance among the donors, there was no evidence of social transfer of resistance, regardless of the ratio of donors and receivers in a group or the time since the pre‐infection. The results suggest that resistance through social signalling may be system‐specific and requires further study into the generality of the phenomenon as well as the nature of the cues involved. peerReviewed

Parasitism is considered a major selective force in natural host populations. Infections can decrease host condition and vigour, and potentially influence, for example, host population dynamics and behavior such as mate choice. We studied parasite infections of two common marine fish species, the sand goby (Pomatoschistus minutus) and the common goby (Pomatoschistus microps), in the brackish water Northern Baltic Sea. We were particularly interested in the occurrence of parasite taxa located in central sensory organs, such as eyes, potentially affecting fish behavior and mate choice. We found that both fish species harbored parasite communities dominated by taxa transmitted to fish through aquatic invertebrates. Infections also showed significant spatiotemporal variation. Trematodes in the eyes were very few in some locations, but infection levels were higher among females than males, suggesting differences in exposure or resistance between the sexes. To test between these hypotheses, we experimentally exposed male and female sand gobies to infection with the eye fluke Diplostomum pseudospathaceum. These trials showed that the fish became readily infected and females had higher parasite numbers, supporting higher susceptibility of females. Eye fluke infections also caused high cataract intensities among the fish in the wild. Our results demonstrate the potential of these parasites to influence host condition and visual abilities, which may have significant implications for survival and mate choice in goby populations.

Even for parasitoids with a wide host range, not all host species are equally suitable, and host qualityoften depends on the plant the host feeds on. We compared oviposition choice and offspring perfor-mance of a generalist pupal parasitoid,Pteromalus apum(Retzius) (Hymenoptera: Pteromalidae), ontwo congeneric hosts reared on two plant species under field and laboratory conditions. The plantscontain defensive iridoid glycosides that are sequestered by the hosts. Sequestration at the pupal stagediffered little between host species and, although the concentrations of iridoid glycosides in the twoplant species differ, there was no effect of diet on the sequestration by host pupae. The rate of success-ful parasitism differed between host species, depending on the conditions they were presented in. Inthe field, where plant-associated cues are present, the parasitoid usedMelitaea cinxia(L.) overMeli-taea athalia(Rottemburg) (Lepidoptera: Nymphalidae), whereas moreM. athaliawere parasitisedin simplified laboratory conditions. In the field, brood size, which is partially determined by rate ofsuperparasitism, depended on both host and plant species. There was little variation in other aspectsof offspring performance related to host or plant species, indicating that the two host plants are ofequal quality for the hosts, and the hosts are of equal quality for the parasitoids. Corresponding tothis, we found no evidence for associative learning by the parasitoid based on their natal host, so withrespect to these host species they are truly generalist in their foraging behaviour. peerReviewed

SUMMARYParasites manipulate their hosts’ phenotype to increase their own fitness. Like any evolutionary adaptation, parasitic manipulations should be costly. Though it is difficult to measure costs of the manipulation directly, they can be evaluated using an indirect approach. For instance, theory suggests that as the parasite infrapopulation grows, the investment of individual parasites in host manipulation decreases, because of cost sharing. Another assumption is that in environments where manipulation does not pay off for the parasite, it can decrease its investment in the manipulation to save resources. We experimentally infected rainbow trout Oncorhynchus mykiss with the immature larvae of the trematode Diplostomum pseudospathaceum, to test these assumptions. Immature D. pseudospathaceum metacercariae are known for their ability to manipulate the behaviour of their host enhancing its anti-predator defenses to avoid concomitant predation. We found that the growth rate of individual parasites in rainbow trout increased with the infrapopulation size (positive density-dependence) suggesting cost sharing. Moreover, parasites adjusted their growth to the intensity of infection within the eye lens where they were localized suggesting population density sensing. Results of this study support the hypothesis that macroparasites can adjust their growth rate and manipulation investment according to cost sharing level and infrapopulation size.

Fish parasites may potentially harm fisheries and aquaculture. Infected fish are unusable, even though they would not pose a direct risk to human health. The aim of this thesis was to investigate the ecology and epidemiology of three previously unknown or poorly known spore-forming parasites infecting muscles of economically important fish species. The first novel microsporidian species, here described as Microsporidium luciopercae, was found from pike-perch (Sander lucioperca) and European perch (Perca fluviatilis) and formed opaque-looking patches in the muscles. The second novel microsporidian species, Myosporidium spraguei, was found from pike-perch and burbot (Lota lota) and occurred within xenomas in the musculature. The third muscle-dwelling parasite, Henneguya zschokkei (Cnidaria), formed white plasmodia in the muscle of whitefish (Coregonus lavaretus). Among the six study lakes, the prevalence of M. luciopercae in wild pike-perch and perch varied from 0 to 1 %, while M. spraguei was more common (prevalence of 5–26 % in pike-perch and 65 % in burbot). M. spraguei was strongly aggregated and most abundant in the middle-sized (37–45 cm) pike-perch and in larger burbot (> 35 cm), but the infection was not related to host sex, sampling season or host condition. In the case of H. zschokkei, in two inland fish farms, one year-old whitefish were virtually uninfected, but the prevalence of infection was higher in 2 and 3 year old fish (up to 36 % of 3 year old fish were infected). New infections appeared seasonally in July-August. Neither the size of plasmodia nor the proportion of different plasmodium types changed from 2 to 3 year old fish, suggesting slow development of H. zschokkei in whitefish. However, the proportion of infected fish with melanised plasmodia increased with fish age, suggesting an age-dependent host response against this strongly aggregated parasite.

Parasites, including macroparasites, protists, fungi, bacteria and viruses, can impose a heavy burden upon host animals. However, hosts are not without defences. One aspect of host defence, behavioural avoidance, has been studied in the terrestrial realm for over 50 years, but was first reported from the aquatic environment approximately 20 years ago. Evidence has mounted on the importance of parasite avoidance behaviours and it is increasingly apparent that there are core similarities in the function and benefit of this defence mechanism between terrestrial and aquatic systems. However, there are also stark differences driven by the unique biotic and abiotic characteristics of terrestrial and aquatic (marine and freshwater) environments. Here, we review avoidance behaviours in a comparative framework and highlight the characteristics of each environment that drive differences in the suite of mechanisms and cues that animals use to avoid parasites. We then explore trade-offs, potential negative effects of avoidance behaviour and the influence of human activities on avoidance behaviours. We conclude that avoidance behaviours are understudied in aquatic environments but can have significant implications for disease ecology and epidemiology, especially considering the accelerating emergence and re-emergence of parasites. This article is part of the Theo Murphy meeting issue ‘Evolution of pathogen and parasite avoidance behaviours'.