Specific features of parasite communities of Siberian cisco Coregonus sardinella in polytypic water bodies of the polar regions: species richness.

The article dwells upon characteristics of the species richness of parasite communities in least cisco Coregonus sardinella Valenciennes in Polar water bodies: Kureyskoye Reservoir, Lake Kutaramakan, Khatanga River. The communities consist of parasite species common in coregonids. The main species in all the parasite communities in least cisco are Dibothriocephalus ditremus and D. dendriticus. Triaenophorus crassus plerocercoids dominate in the regulated water body (Kureyskoye Reservoir). To analyze the species richness of the component communities (s), coefficients of the power-law equation s = a × nb were applied, where a is the expected species richness in the first host individual examined, and b is the rate of species accumulation in parasite component communities (s). The resulting curves of species accumulation give statistical rigor to the assessment of differences and similarities of the species richness of the component communities of parasites in least cisco of various ages and habitats. The species richness of infra- and component parasite communities in least cisco was found to be significantly lower in the lake and the storage reservoir than in riverine habitats (Khatanga River). At the same time, the species accumulation rate for parasite communities in least cisco from Lake Kutaramakan was higher than in the reservoir due to a higher diversity of parasite infracommunities.

A report on the complete mitochondrial genome of the trematode Azygia robusta Odhner, 1911, its new definitive host from the Russian Far East, and unexpected phylogeny of Azygiidae within Digenea, as inferred from mitogenome sequences.

New data on the complete mitochondrial genome of Azygia robusta (Azygiidae) were obtained by the next-generation sequencing (NGS) approach. The mitochondrial DNA (mtDNA) of A. robusta had a length of 13 857 bp and included 12 protein-coding genes, two ribosomal genes, 22 transfer RNA genes, and two non-coding regions. The nucleotide sequences of the complete mitochondrial genomes of two A. robusta specimens differed from each other by 0.12 ± 0.03%. Six of 12 protein-coding genes demonstrated intraspecific variation. The difference between the nucleotide sequences of the complete mitochondrial genomes of A. robusta and Azygia hwangtsiyui was 26.95 ± 0.35%; the interspecific variation of protein-coding genes between A. robusta and A. hwangtsiyui ranged from 20.5 ± 0.9% (cox1) to 30.7 ± 1.2% (nad5). The observed gene arrangement in the mtDNA sequence of A. robusta was identical to that of A. hwangtsiyui. Codon usage and amino acid frequencies were highly similar between A. robusta and A. hwangtsiyui. The results of phylogenetic analyses based on mtDNA protein-coding regions showed that A. robusta is closely related to A. hwangtsiyui (belonging to the same suborder, Azygiida) that formed a distinct early-diverging branch relative to all other Digenea. A preliminary morphological analysis of paratypes of the two azygiid specimens studied showed visible morphological differences between them. The specimen extracted from Sakhalin taimen (Parahucho perryi) was most similar to A. robusta. Thus, we here provide the first record of a new definitive host, P. perryi, for A. robusta and also molecular characteristics of the trematode specimens.

Efficacy of hydrogen peroxide to reduce Gyrodactylus species infestation density on four fish species.

OBJECTIVE: The ability to effectively treat parasitic infestations of fish is of high importance for fish culture facilities. However, tools or approved therapies for treating infestations on fish are limited. This paper summarizes results from four separate clinical field studies that evaluated the efficacy of hydrogen peroxide (H2 O2 ; 35% PEROX-AID) for reducing Gyrodactylus spp. infestation density. METHODS: Three species of Gyrodactylus were studied (G. salmonis, hosts: Brook Trout Salvelinus fontinalis and Lake Trout S. namaycush; G. freemani, host: Yellow Perch Perca flavescens; G. hoffmani, host: Fathead Minnow Pimephales promelas) before and after the application of immersion H2 O2 therapy. RESULT: Parasite density was significantly reduced for each parasite × host combination to which H2 O2 therapy was applied. Two clinical field studies in salmonids were found to demonstrate substantial effectiveness that enabled 35% PEROX-AID approval. CONCLUSION: Further assessments of Gyrodactylus spp. could expand the use of H2 O2 for controlling these parasites in aquaculture. Specifically, H2 O2 was effective at all levels tested (50 or 75 mg H2 O2 /L for 60 min for the Yellow Perch and Fathead Minnow clinical field studies; 100 or 150 mg H2 O2 /L for 30 min regardless of salt pre-treatment for the Brook Trout study; and 100 mg H2 O2 /L for 30 min or 50 mg H2 O2 /L for 60 min for the Lake Trout study).

Mining various genomic resources to resolve old alpha-taxonomy questions: A test of the species hypothesis of the Proteocephalus longicollis species complex (Cestoda: Platyhelminthes) from salmonid fishes.

High-throughput sequencing strategies became commonly employed to study non-model parasites, but the corresponding genomes and transcriptomes were seldom mined following the original publication. Similar to the data generated with genome skimming techniques based on shallow-depth shotgun genomes, various genomic and transcriptomic resources can be screened for useful molecular phylogenetic markers traditionally characterised with Sanger sequencing. Here, we provide an example of a strategy using reduced-representation genomic as well as transcriptomic data to obtain broad insights into the molecular diversity of the cestode Proteocephalus longicollis, a common parasite of salmonids distributed throughout the Holarctic region. We extract popular mitochondrial and nuclear ribosomal markers from various genomic resources for hundreds of parasite specimens from multiple European whitefish populations and compare those with Proteocephalus representatives from other species of salmonids and various geographical regions. In contrast with the previous morphology-based assessments, molecular phylogeny reveals a high degree of genetic divergence between Proteocephalus isolates from different salmonids, contrastingly low genetic differentiation within the parasite's populations hosted by the European whitefish (Coregonus lavaretus species complex), and a sister species relationship of Proteocephalus from European whitefish and Proteocephalus percae, a parasite of European perch (Perca fluviatilis). Proteocephalus spp. from North American lake whitefish, brown trout and Arctic charr each formed clearly distinct lineages. These results advance our understanding of the interrelationships of the Proteocephalus-aggregate, a well-recognized clade of Holarctic freshwater fish proteocephalids, and support resurrection of some of the nominal species of Proteocephalus, including Proteocephalus exiguus La Rue, 1911 from North American coregonids and Proteocephalus fallax La Rue, 1911 from European C. lavaretus, reserving Proteocephalus longicollis (Zeder, 1800) exclusively for parasites of Salmo trutta.

Using a mechanistic framework to model the density of an aquatic parasite Ceratonova shasta.

Ceratonova shasta is a myxozoan parasite endemic to the Pacific Northwest of North America that is linked to low survival rates of juvenile salmonids in some watersheds such as the Klamath River basin. The density of C. shasta actinospores in the water column is typically highest in the spring (March-June), and directly influences infection rates for outmigrating juvenile salmonids. Current management approaches require quantities of C. shasta density to assess disease risk and estimate survival of juvenile salmonids. Therefore, we developed a model to simulate the density of waterborne C. shasta actinospores using a mechanistic framework based on abiotic drivers and informed by empirical data. The model quantified factors that describe the key features of parasite abundance during the period of juvenile salmon outmigration, including the week of initial detection (onset), seasonal pattern of spore density, and peak density of C. shasta. Spore onset was simulated by a bio-physical degree-day model using the timing of adult salmon spawning and accumulation of thermal units for parasite development. Normalized spore density was simulated by a quadratic regression model based on a parabolic thermal response with river water temperature. Peak spore density was simulated based on retained explanatory variables in a generalized linear model that included the prevalence of infection in hatchery-origin Chinook juveniles the previous year and the occurrence of flushing flows (≥171 m3/s). The final model performed well, closely matched the initial detections (onset) of spores, and explained inter-annual variations for most water years. Our C. shasta model has direct applications as a management tool to assess the impact of proposed flow regimes on the parasite, and it can be used for projecting the effects of alternative water management scenarios on disease-induced mortality of juvenile salmonids such as with an altered water temperature regime or with dam removal.

Potential negative effects and heterogeneous distribution of a parasitic copepod Salmincola edwardsii (Copepoda: Lernaeopodidae) on Southern Asian Dolly Varden Salvelinus curilus in Hokkaido, Japan.

The genus Salmincola is an ectoparasitic copepod group commonly infesting the branchial and buccal cavities of salmonids. While negative impacts on hatchery fishes have been reported, their impacts on wild fish populations and distribution patterns are critically understudied. In the Shiretoko Peninsula, Hokkaido, Japan, we found parasites belonging to this genus on the branchial cavity of a stream salmonid, Southern Asian Dolly Varden Salvelinus curilus. All parasites recovered were identified as Salmincola edwardsii based on morphological characteristics and partial 28S rDNA sequences. Prevalence was highly heterogeneous even among neighboring streams (0-54.8%, < 10 km) with the mean intensity among streams being generally low (2.19 parasites/infeted fish). Despite the low intensity, quantile regression analysis showed negative trends between parasite intensity and host condition, suggesting that the infestation of S. edwardsii has a potential negative impact on the host salmonid. In addition, a single copepod was found from an anadromous fish, which could indicate some salinity tolerance of the copepods. It is important to evaluate the effects of Salmincola spp. on host species and determine the limiting factors on the parasite's distribution for proper management.

Simulated effects of increasing salmonid production on sea lice populations in Norway.

Norway produces more than one million tonnes of salmonids every year, almost exclusively in open-water net pens. In 2014, the Norwegian government announced plans to increase salmonid production. However, increasing the number of farmed salmonids can have negative effects on the marine environment that threaten the industry's sustainability. In particular, production growth can lead to an increase in density-dependent diseases, including parasitic sea lice. The aim of this study was to simulate the effects of increased salmonid production on sea lice abundance using different scenarios for increasing the number of fish and for the management of sea lice. We used a previously developed, partly stage-structured model based on Norwegian production and environmental data to simulate the different scenarios. Our results show that increasing the marine farmed salmonid population at a national level by two or five times the current production leads to an increase in the sea lice abundance by 3.5% and 7.1%, respectively. We also found that by lowering the maximum allowable level of sea lice to an average of 0.049 adult females per fish, weekly treatments can be used to control sea lice population growth with a five times increase in production. However, this increases the number of farms treating per week by as much as 281.3%, which can lead to high costs and increased mortality among farmed salmonids. Overall, the results from our study shed light on the effects of increasing salmonid production in Norway with respect to the ongoing threat of sea lice infestations.

Feeding habits shape infection levels by plerocercoids of the tapeworm Triaenophorus crassus in muscle of a sympatric pair of whitefish in an oligotrophic lake.

Lake Teletskoye (West Siberia, Russia) is inhabited by a sympatric pair of whitefish, with each member of the pair being characterized by different feeding habits. Coregonus lavaretus pidschian (Gmelin, 1789) is a large 'benthivorous' form, while C. l. pravdinellus (Dulkeit, 1949) is a small 'planktivorous' form. Fish were collected from the end of August to the middle of September in 2017 and 2019-2020 in the north part of Lake Teletskoye. For the 'benthivorous' form the prevalence, intensity and abundance of T. crassus ranged from 22.4% to 51.9%, 1.9-2.8 and 0.4-1.3, respectively, whereas the same indices for the 'planktivorous' form ranged from 94.7% to 97.5%, 4.2-4.8 and 4.0-4.7, respectively. The level of prevalence of infection and abundance of T. crassus in muscle was relatively stable among studied years for both forms. The level of prevalence was higher in the years 2019 and 2020 than in 2017 for the 'benthivorous' form, whereas for the 'planktivorous' form this index did not change during the studied years. For the first time, a partial sequencing of the cox1 gene (593 bp) for T. crassus was sequenced. All 15 plerocercoids of T. crassus were represented by four haplotypes.

Ultrastructural observations on the oncomiracidium epidermis and adult tegument of Discocotyle sagittata, a monogenean gill parasite of salmonids.

During their different life stages, parasites undergo remarkable morphological, physiological, and behavioral "metamorphoses" to meet the needs of their changing habitats. This is even true for ectoparasites, such as the monogeneans, which typically have a free-swimming larval stage (oncomiracidium) that seeks out and attaches to the external surfaces of fish where they mature. Before any obvious changes occur, there are ultrastructural differences in the oncomiracidium's outer surface that prepare it for a parasitic existence. The present findings suggest a distinct variation in timing of the switch from oncomiracidia epidermis to the syncytial structure of the adult tegument and so, to date, there are three such categories within the Monogenea: (1) Nuclei of both ciliated cells and interciliary cytoplasm are shed from the surface layer and the epidermis becomes a syncytial layer during the later stages of embryogenesis; (2) nuclei of both ciliated cells and interciliary syncytium remain distinct and the switch occurs later after the oncomiracidia hatch (as in the present study); and (3) the nuclei remain distinct in the ciliated epidermis but those of the interciliary epidermis are lost during embryonic development. Here we describe how the epidermis of the oncomiracidium of Discocotyle sagittata is differentiated into two regions, a ciliated cell layer and an interciliary, syncytial cytoplasm, both of which are nucleated. The interciliary syncytium extends in-between and underneath the ciliated cells and sometimes covers part of their apical surfaces, possibly the start of their shedding process. The presence of membranous whorls and pyknotic nuclei over the surface are indicative of membrane turnover suggesting that the switch in epidermis morphology is already initiated at this stage. The body tegument and associated putative sensory receptors of subadult and adult D. sagittata are similar to those in other monogeneans.

Hatching and survival of the salmon 'gill maggot' Salmincola californiensis (Copepoda: Lernaeopodidae) reveals thermal dependence and undocumented naupliar stage.

Salmincola californiensis is a Lernaeopodid copepod parasitizing Pacific salmon and trout of the genus Oncorhynchus. Salmincola californiensis is of increasing concern in both native and introduced ranges because of its potential fish health impacts and high infection prevalence and intensity in some systems. Discrepancies in the documented life history phenology of S. californiensis with the sister species Salmincola edwardsii, as well as our laboratory observations, led us to question the existing literature. We documented a naupliar stage, thought lost for S. californiensis. In addition, we found a high degree of thermal sensitivity in egg development, with eggs developing faster under warmer conditions. Survival of copepodids was also highly dependent on temperature, with warmer conditions reducing lifespan. The longest lived copepodid survived 18 days at 4°C in stark contrast to the generally accepted <48 h survival for that life stage. We also note a consistent relationship between egg sac size and the number of eggs contained. However, egg sac sizes were highly variable. Our findings demonstrate that revisiting old assumptions for S. californiensis and related taxa will be a necessary step to improving our knowledge of the parasite life history and development that will be critical to disease management.

In vitro and in vivo assays reveal that cations affect nematocyst discharge in Myxobolus cerebralis (Cnidaria: Myxozoa).

Myxozoans are parasitic, microscopic cnidarians that have retained the phylum-characteristic stinging capsules called nematocysts. Free-living cnidarians, like jellyfish and corals, utilize nematocysts for feeding and defence, with discharge powered by osmotic energy. Myxozoans use nematocysts to anchor to their fish hosts in the first step of infection, however, the discharge mechanism is poorly understood. We used Myxobolus cerebralis, a pathogenic myxozoan parasite of salmonid fishes, and developed two assays to explore the nature of its nematocyst discharge. Using parasite actinospores, the infectious stage to fish, we stimulated discharge of the nematocysts with rainbow trout mucus in vitro, in solutions enriched with chloride salts of Na+, K+, Ca2+ and Gd3+, and quantified discharge using microscopy. We then used quantitative polymerase chain reaction to evaluate the in vivo effects of these treatments, plus Mg2+ and the common aquaculture disinfectant KMnO4, on the ability of M. cerebralis actinospores to infect fish. We found that Mg2+ and Gd3+ reduced infection in vivo, whereas Na+ and K+ over-stimulated nematocyst discharge in vitro and reduced infection in vivo. These findings align with nematocyst discharge behaviour in free-living Cnidaria, and suggest phylum-wide commonalties, which could be exploited to develop novel approaches for controlling myxozoan diseases in aquaculture.

Morphological and molecular data show no evidence of the proposed replacement of endemic Pomphorhynchus tereticollis by invasive P. laevis in salmonids in southern Germany.

Changes in parasite communities might result in new host-parasite dynamics and may threaten local fish populations. This phenomenon has been suggested for acanthocephalan parasites in the river Rhine and Danube where the species Pomphorhynchus tereticollis is becoming replaced by the Ponto-Caspian P. laevis. Developing knowledge on morphologic, genetic and behavioural differences between such species is important to follow such changes. However, disagreements on the current phylogeny of these two acanthocephalan species are producing conflicts that is affecting their correct identification. This study is offering a clearer morphological and genetic distinction between these two species. As P. tereticollis is found in rhithral tributaries of the Rhine, it was questioned whether the local salmonid populations were hosts for this species and whether P. laevis was expanding into the Rhine watershed as well. In order to test for this, brown trout, Salmo trutta, and grayling, Thymallus thymallus from South-Western Germany watersheds have been samples and screened for the occurrence of acanthocephalan parasites. For the first time, both species were confirmed to be hosts for P. tereticollis in continental Europe. P. tereticollis was found to be common, whereas P. leavis was found only at a single location in the Danube. This pattern suggest either that the expansion of P. laevis through salmonid hosts into rhithral rivers has not yet occurred, or that not yet ascertained biotic or abiotic features of rhithral rivers hinder P. laevis to spread into these areas.

Unexpected diversity in northern Europe: trematodes from salmonid fishes in Iceland with two new species of Crepidostomum Braun, 1900.

In 2018 and 2019, salmonid fishes, Salmo trutta L. and Salvelinus alpinus (L.) from lakes in Iceland were assessed for trematodes during a parasitological examination. Combined morphological and molecular analyses revealed the presence of four trematode species, two of which were previously known to parasitise salmonids in Iceland, Crepidostomum farionis (Müller, 1780) and Phyllodistomum umblae (Fabricius, 1780), and the two remaining species were recognised as new to science. Morphologically and genetically, Crepidostomum brinkmanni n. sp. and C. pseudofarionis n. sp. are closely related to two European species, namely C. metoecus and C. farionis. However, C. brinkmanni n. sp. is distinct by the position of maximum body width and arrangement of vitelline follicles; C. pseudofarionis n. sp. is distinct by its stout body, position of maximum body width, size of muscular papillae relative to oral sucker and the anterior extent of vitelline follicles. The new species were previously molecularly detected in their intermediate and definitive hosts in Norway and Ukraine, but their sequences were not supplemented with any morphological characterisation. In the present study, we provide detailed morphological descriptions and molecular sequences (28S rDNA and ITS2) of the four species of trematodes detected in Iceland. The discovery of the two new species of Crepidostomum indicates that the trematode diversity in fishes in the north is higher than previously known; our finding doubles the species spectrum of fish trematodes for Iceland. The record of C. brinkmanni from Ukraine indicates that its distribution might not be limited to northern latitudes.

Proliferative kidney disease in Alaskan salmonids with evidence that pathogenic myxozoans may be emerging north.

Proliferative kidney disease (PKD) of salmonids, a chronic immunopathology caused by the myxozoan parasite Tetracapsuloides bryosalmonae, is exacerbated by increased water temperatures. PKD causes economic concerns to trout farmers and contributes to the decline of wild salmonid populations in North America and Europe. The parasite occurs as far north as Norway and Iceland in Europe and was confirmed from California to southern British Columbia in the American continent. In mid-September 2011 adult chum salmon (Oncorhynchus keta) were sampled from Kantishna River, a tributary to Yukon River in Alaska. Clinical PKD was diagnosed based on the macroscopic appearance of mottled kidneys that were uniformly swollen and by the detection of tumultuous histozoic extrasporogonic and coelozoic sporogonic stages of T. bryosalmonae in renal tissue by histopathology. Archived samples provided the molecular confirmation and local strain identification, representing the first confirmed case of PKD in wild adult chum salmon, also co-infected with Parvicapsula minibicornis that represents another novel myxozoan detection in Alaska. Our investigation was extended to another case from August/September 1997, with mortality following furunculosis and ectoparasite co-infections, in sockeye salmon (Oncorhynchus nerka) pre-smolts net-pen reared in English Bay Lakes, Alaska. Immunohistochemistry on archived histological preparations confirmed T. bryosalmonae sporogonic and extrasporogonic stages, indicating a severe to resolving PKD, with concomitant Chloromyxum spp. infection. Those cases provide the first documentation that this parasite is present in Alaska and causes PKD in wild and cultured salmonids in the region. The known geographic range of T. bryosalmonae can be extended to ~267 km south of the Arctic Circle, representing the northernmost detection in America. Given the vast size of Alaska and small resident population, it is likely that T. bryosalmonae remained undetected, but more recently became evident due to the clinical manifestation of PKD, possibly linked to increasing water temperatures reported at the sample locations.

Redescription of Rhadinorhynchus trachuri Harada, 1935 (Acanthocephala: Rhadinorhynchidae) from Marine Fish in Vietnam and California with a Discussion of its Zoogeography.

BACKGROUND: Rhadinorhynchus trachuri Harada, 1935 is probably one of the most widely distributed acanthocephalans in the world, but the least known taxonomically. This report fills the large gap of knowledge about its taxonomy, host relationships, and worldwide distribution in marine fishes of both sides of the Pacific Ocean and the connecting Indian Ocean. PURPOSE: To revisit the taxonomy of this species, provide a more complete morphological description and expand our knowledge about its morphological identity and variations and elaborate on its worldwide zoogeography; accounting for its worldwide distribution based on new collections, museum specimens, and on published reports. METHODS: Specimens from Vietnam were processed, identified, and documented using standard techniques. Specimens of R. trachuri were studied from three species of fish in three families, Auxis thazard (Lacépède) (Scombridae), Megalaspis cordyla (Linn.) (Carangidae), and Tylosurus sp. (Belonidae) in Nha Trang and Binh Thuan in the south Pacific coast of Vietnam and additional museum specimens mostly from salmonids from the Pacific coast of California were also studied. Extensive research was conducted on published records of R. trachuri and its marine fish hosts worldwide. RESULTS: Specimens of R. trachuri are characterized by cylindrical trunk with anterior rings of spines and posterior ventral and lateral spines, a proboscis usually with 12 hook rows each with 19-24 hooks, variable position of testes, and a long uterus, uterine bell attached to the dorsal trunk wall, rounded vagina, and unusually dorsal gonopore. The hosts and geographical distribution of R. trachuri suggest a northern linkage of infections between the Asian and American Pacific populations across the Bering Sea involving salmonid fishes and a southern linkage through the Indian Ocean involving Carangid fishes. Other hosts involved in the cycling and distribution of infections are also discussed with some definitive hosts also serving as paratenic hosts. CONCLUSIONS: This is the first complete description about R. trachuri and the first comprehensive accounting of its worldwide distribution in 19 species of marine fish in seven families. Considerable intraspecific variability related to geographical restrictions, intermediate and definitive host specificity and distribution, and host feeding behavior was documented.

Evolutionary dynamics of Ceratonova species (Cnidaria: Myxozoa) reveal different host adaptation strategies.

The myxozoan parasite Ceratonova shasta is an important pathogen that infects multiple species of Pacific salmonids. Ongoing genetic surveillance has revealed stable host-parasite relationships throughout the parasite's endemic range. We applied Bayesian phylogenetics to test specific hypotheses about the evolution of these host-parasite relationships within the well-studied Klamath River watershed in Oregon and California, USA. The results provide statistical support that different genotypes of C. shasta are distinct lineages of one species, which is related to two other Ceratonova species in the same ecosystems; Ceratonova X in speckled dace and C. gasterostea in threespine stickleback. Furthermore, we found strong support for the hypothesis that C. shasta type 0 in native steelhead trout and type I in Chinook salmon each evolved with a specialist host adaptation strategy, while C. shasta type II in coho salmon resulted from a generalist host adaptation strategy. Inferred date and host species of the most recent common ancestor of extant Klamath basin types indicate that it occurred between 14,000 and 21,000 years ago, and most likely infected a native steelhead or rainbow trout host.

Host Switching of Zoonotic Broad Fish Tapeworm (Dibothriocephalus latus) to Salmonids, Patagonia.

Diphyllobothriosis is a reemerging zoonotic disease because of global trade and increased popularity of eating raw fish. We present molecular evidence of host switching of a human-infecting broad fish tapeworm, Dibothriocephalus latus, and use of salmonids as intermediate or paratenic hosts and thus a source of human infection in South America.

A partly stage-structured model for the abundance of salmon lice in salmonid farms.

The parasitic salmon louse constrains growth in the Norwegian salmon farming industry through density dependent host-parasite interaction. Hence, there is a need for insight into how increases in salmon production, varying spatial organisation of the production and pest control strategies affect salmon louse population dynamics. Here we present a new salmon louse model for exploring effects of varying salmon farming conditions on spatio-temporal abundances of the parasite. The salmon louse model is partly stage-structured, comprising of (i) adult female lice and (ii) other mobile stages of lice. The abundance of adult females depend on survival of females from previous weeks and recruitment from the other mobile group of lice. The other mobiles also depend on survival of other mobiles from previous weeks, as well as recruitment from the previous generation of reproducing adult females from the same farm or from farms in the neighbourhood. In addition, expected abundances of the two stage-groups are modelled as functions of biological and physical covariates. The model is fitted to weekly salmon farm data covering all marine farms producing salmonids along the Norwegian cost over the years 2012-2016. Among novel results from fitting the model are estimates of the time-lag structure representing recruitment of other mobile lice from the previous generation adult females for different temperatures. Furthermore, the model estimates how various factors affect the susceptibility of fish on farms to louse infection and effects of treatments to control infection. Finally, the model estimates density dependent effects of increasing the number of fish in farms and of increasing the numbers of farms, on the rate of recruitment of other mobile lice. Analytically, the parameters representing density dependencies suggest that few farms with many fish and large between farm distances is effective in terms of minimising the overall output of salmon lice infection.

A RNAi-based therapeutic proof of concept targets salmonid whirling disease in vivo.

Myxobolus cerebralis is a cnidarian-myxozoan parasite that causes salmonid whirling disease. M. cerebralis alternates between two hosts: (1) a vertebrate salmonid and (2) an invertebrate oligochaete, Tubifex tubifex. There is no successful treatment for salmonid whirling disease. MyxSP-1 is a M. cerebralis serine protease implicated in whirling disease pathogenesis. We hypothesized that short-interfering RNA (siRNA)-induced RNA interference (RNAi) can silence MyxSP-1 in the invertebrate host and abrogate the M. cerebralis life cycle. This would preclude whirling disease infection in the salmonid host. To test this hypothesis, we first developed a siRNA delivery protocol in T. tubifex. Second, we determined the effective dose for siRNA treatment of M. cerebralis-infected T. tubifex. M. cerebralis-infected T. tubifex were treated with different concentrations of MyxSP-1 or negative control siRNAs (1µM, 2µM, 5µM or 7µM) at 15°C for 24h, 48h, 72h and 96h, respectively. We monitored MyxSP-1 knockdown using real-time quantitative PCR (qPCR). siRNA treatment with MyxSP-1 siRNA at 2µM concentration for 24h at 15°C showed maximum significant MyxSP-1 knockdown in T. tubifex. Third, we determined the time points in the M. cerebralis life cycle in T. tubifex at which siRNA treatment was most effective. M. cerebralis-infected T. tubifex were treated with MyxSP-1 or negative control siRNAs (2µM concentration for 24h at 15°C) at 24 hours post-infection (24hpi), 48hpi, 72hpi, 96hpi, 1 month post-infection (1mpi), 2mpi and 3mpi, respectively. We observed that siRNA treatment of T. tubifex was most effective at 1mpi, 2mpi and 3mpi. Fourth, we immersed specific-pathogen-free rainbow trout fry in water inhabited by MyxSP-1 siRNA-treated T. tubifex (at 1mpi, 2mpi and 3mpi). The salmonids did not develop whirling disease and showed significant MyxSP-1 knockdown. We also observed long-term RNAi in T. tubifex. Together these results demonstrate a novel RNAi-based therapeutic proof of concept in vivo against salmonid whirling disease.

Learned parasite avoidance is driven by host personality and resistance to infection in a fish-trematode interaction.

Cognitive abilities related to the assessment of risk improve survival. While earlier studies have examined the ability of animals to learn to avoid predators, learned parasite avoidance has received little interest. In a series of behavioural trials with the trematode parasite Diplostomum pseudospathaceum, we asked whether sea trout (Salmo trutta trutta) hosts show associative learning in the context of parasitism and if so, whether learning capacity is related to the likelihood of infection mediated through host personality and resistance. We show that animals are capable of learning to avoid visual cues associated with the presence of parasites. However, avoidance behaviour ceased after the likely activation of host resistance following consecutive exposures during learning, suggesting that resistance to infection outweighs avoidance. Further, we found a positive relationship between learning ability and boldness, suggesting a compensation of risky lifestyles through increased investment in cognitive abilities. By contrast, an increased risk of infection due to low resistance was not balanced by learning ability. Instead, these traits were positively related, which may be explained by inherent physiological qualities controlling both traits. Overall, the results demonstrate that parasitism, in addition to other biological interactions such as predation, is an important selective factor in the evolution of animal cognition.