Non-lethal detection of Eubothrium crassum (Cestoda) in farmed Atlantic salmon, Salmo salar, using anal swabs and real-time PCR.

Detection of intestinal parasites in fish typically requires autopsy, resulting in the sacrifice of the fish. Here, we describe a non-lethal method for detecting the tapeworm Eubothrium crassum in fish using anal swabs and real-time PCR detection. Two assays were developed to detect cytochrome oxidase I (COI) mitochondrial DNA and 18S ribosomal DNA sequences of E. crassum, respectively. The assays were tested on swab samples from confirmed pathogen free Atlantic salmon (Salmo salar L.) and on samples from farmed Atlantic salmon, where the presence and intensity of parasites had been established through autopsy. The COI assay was shown to be specific to E. crassum, while the 18S assay also amplified the closely related E. salvelini, a species infecting Arctic charr (Salvelinus alpinus L.) in freshwater. The COI assay detected E. crassum in all field samples regardless of parasite load while the 18S assay failed to detect the parasite in two samples. The results thus demonstrates that this non-lethal approach can effectively detect E. crassum and can be a valuable tool in assessing the prevalence of infection in farmed salmon, aiding in treatment decisions and evaluating treatment effectiveness.

Spatial and intra-host distribution of myxozoan parasite Tetracapsuloides bryosalmonae among Baltic sea trout (Salmo trutta).

Proliferative kidney disease caused by the myxozoan parasite Tetracapsuloides bryosalmonae has been actively studied in juvenile salmonids for decades. However, very little is known about parasite prevalence and its geographical and intra-host distribution at older life stages. We screened T. bryosalmonae among adult sea trout (Salmo trutta) (n = 295) collected along the Estonian Baltic Sea coastline together with juvenile trout from 33 coastal rivers (n = 1752) to assess spatial infection patterns of the adult and juvenile fish. The parasite was detected among 38.6% of adult sea trout with the prevalence increasing from west to east, and south to north, along the coastline. A similar pattern was observed in juvenile trout. Infected sea trout were also older than uninfected fish and the parasite was detected in sea trout up to the age of 6 years. Analysis of intra-host distribution of the parasite and strontium to calcium ratios from the otoliths revealed that (re)infection through freshwater migration may occur among adult sea trout. The results of this study indicate that T. bryosalmonae can persist in a brackish water environment for several years and that returning sea trout spawners most likely contribute to the parasite life cycle by transmitting infective spores.

Evidence for infection influencing survival of the freshwater copepod Salmincola californiensis, a parasite of Pacific salmon and trout.

OBJECTIVE: We explore apparent infection of Salmincola californiensis arising during investigations involving this lernaeopodid copepod parasitic on Pacific salmon and trout Oncorhynchus spp. METHODS: We noted occasional unusual coloration of adult female copepods collected from the wild. These females were bright blue and pink in contrast to the cream white coloration characteristic of the copepod. We also observed that similar color patterns developed under laboratory settings when copepod eggs were held for hatching. In paired egg cases, we found consistent hatching failure of blue and pink eggs and patterns in apparent disease development that would be consistent with both vertical and horizontal transmission. RESULT: Attempts to identify the cause of the apparent infection using genetic methods and transmission electron microscopy were inconclusive. CONCLUSION: Iridovirus infection was initially suspected, but bacterial infection is also plausible. This apparent reduced hatching success of S. californiensis warrants further exploration as it could reduce local abundances. Given the potential importance of a disease impacting this copepod, a parasite that itself affects endangered and commercially important Pacific salmon and trout, future research would benefit from clarification of the apparent infection through additional sequencing, primer development, visualization, and exploration into specificity and transmission.

Genome-wide alternative splicing profile in the posterior kidney of brown trout (Salmo trutta) during proliferative kidney disease.

BACKGROUND: The cnidarian myxozoan parasite Tetracapsuloides bryosalmonae causes chronic proliferative kidney disease (PKD) in salmonids. This parasite is a serious threat to wild and cultured salmonids. T. bryosalmonae undergoes intra-luminal sporogonic development in the kidney of brown trout (Salmo trutta) and the viable spores are released via urine. We investigated the alternative splicing pattern in the posterior kidney of brown trout during PKD. RESULTS: RNA-seq data were generated from the posterior kidney of brown trout collected at 12 weeks post-exposure to T. bryosalmonae. Subsequently, this data was mapped to the brown trout genome. About 153 significant differently expressed alternatively spliced (DEAS) genes, (delta PSI = 5%, FDR P-value < 0.05) were identified from 19,722 alternatively spliced events. Among the DEAS genes, the least and most abundant alternative splicing types were alternative 5' splice site (5.23%) and exon skipping (70.59%), respectively. The DEAS genes were significantly enriched for sodium-potassium transporter activity and ion homeostasis (ahcyl1, atp1a3a, atp1a1a.1, and atp1a1a.5). The protein-protein interaction network analysis enriched two local network clusters namely cation transporting ATPase C-terminus and Sodium/potassium ATPase beta chain cluster, and mixed inclusion of Ion homeostasis and EF-hand domain cluster. Furthermore, the human disease-related salmonella infection pathway was significantly enriched in the protein-protein interaction network. CONCLUSION: This study provides the first baseline information about alternative splicing in brown trout during PKD. The generated data lay a foundation for further functional molecular studies in PKD - brown trout infection model. The information generated from the present study can help to develop therapeutic strategies for PKD in the future.

Mitigating human impacts including climate change on proliferative kidney disease in salmonids of running waters.

Over the last two decades, an increasing number of reports have identified a decline in salmonid populations, possibly linked to infection with the parasite Tetracapsuloides bryosalmonae and the corresponding disease, that is, proliferative kidney disease (PKD). The life cycle of this myxozoan parasite includes sessile bryozoan species as invertebrate host, which facilitates the distribution of the parasite in running waters. As the disease outcome is temperature dependent, the impact of the disease on salmonid populations is increasing with global warming due to climate change. The goal of this review is to provide a detailed overview of measures to mitigate the effects of PKD on salmonid populations. It first summarizes the parasite life cycle, temperature-driven disease dynamics and new immunological and molecular research into disease resistance and, based on this, discusses management possibilities. Sophisticated management actions focusing on local adaptation of salmonid populations, restoration of the riverine ecosystem and keeping water temperatures cool are necessary to reduce the negative effects of PKD. Such actions include temporary stocking with PKD-resistant salmonids, as this may assist in conserving current populations that fail to reproduce.

Know your enemy - transcriptome of myxozoan Tetracapsuloides bryosalmonae reveals potential drug targets against proliferative kidney disease in salmonids.

The myxozoan Tetracapsuloides bryosalmonae is a widely spread endoparasite that causes proliferative kidney disease (PKD) in salmonid fish. We developed an in silico pipeline to separate transcripts of T. bryosalmonae from the kidney tissue of its natural vertebrate host, brown trout (Salmo trutta). After stringent filtering, we constructed a partial transcriptome assembly T. bryosalmonae, comprising 3427 transcripts. Based on homology-restricted searches of the assembled parasite transcriptome and Atlantic salmon (Salmo salar) proteome, we identified four protein targets (Endoglycoceramidase, Legumain-like protease, Carbonic anhydrase 2, Pancreatic lipase-related protein 2) for the development of anti-parasitic drugs against T. bryosalmonae. Earlier work of these proteins on parasitic protists and helminths suggests that the identified anti-parasitic drug targets represent promising chemotherapeutic candidates also against T. bryosalmonae, and strengthen the view that the known inhibitors can be effective in evolutionarily distant organisms. In addition, we identified differentially expressed T. bryosalmonae genes between moderately and severely infected fish, indicating an increased abundance of T. bryosalmonae sporogonic stages in fish with low parasite load. In conclusion, this study paves the way for future genomic research in T. bryosalmonae and represents an important step towards the development of effective drugs against PKD.

Higher mortality of the less suitable brown trout host compared to the principal Atlantic salmon host when infested with freshwater pearl mussel (Margaritifera margaritifera) glochidia.

The freshwater pearl mussel (Margaritifera margaritifera) is a highly host-specific parasite, with an obligate parasitic stage on salmonid fish. Atlantic salmon (Salmo salar) and brown trout (Salmo trutta f. trutta and Salmo trutta f. fario) are the only hosts in their European distribution. Some M. margaritifera populations exclusively infest either Atlantic salmon or brown trout, while others infest both hosts with one salmonid species typically being the principal host and the other a less suitable host. Glochidial abundance, prevalence and growth are often used as parameters to measure host suitability, with the most suitable host species displaying the highest parameters. However, it is not known if the degree of host specialisation will negatively influence host fitness (virulence) among different host species. In this study we examined the hypothesis that glochidial infestation would result in differential virulence in two salmonid host species and that lower virulence would be observed on the most suitable host. Atlantic salmon and brown trout were infested with glochidia from two M. margaritifera populations that use Atlantic salmon as their principal host, and the difference in host mortality among infested and control (sham infested) fish was examined. Higher mortality was observed in infested brown trout (the less suitable host) groups, compared to the other test groups. Genetic assignment was used to identify offspring from individual mother mussels. We found that glochidia from individual mothers can infest both the salmonid hosts; however, some mothers displayed a bias towards either salmon or trout. We believe that the differences in host-dependent virulence and the host bias displayed by individual mothers were a result of genotype × genotype interactions between the glochidia and their hosts, indicating that there is an underlying genetic component for this parasite-host interaction.

Morphological and molecular characterization of a novel Myxobolus species from the gastrointestinal tract of brown trout (Salmo trutta) in Spain.

The genus Myxobolus Bütschli, 1882 is the largest group within the class Myxosporea and includes 905 nominal species, 18 of which have been found to infect fish belonging to the family Salmonidae. In the present study, microscopic analysis enabled detection of myxospores in 43 of 613 (7.0%) gastrointestinal tracts from brown trout (Salmo trutta) captured in several rivers in the northwest of the Iberian Peninsula. Measurement of the whole myxospores, polar capsules and other morphological characteristics, together with identification of the site of infection, has led us to propose a novel salmonid-myxobolid species, Myxobolus compostellanus n. sp. Molecular analysis of the small subunit ribosomal RNA (SSU-rRNA) gene yielded the same consensus sequence of 2039 bp in 14 fish specimens. A BLAST search indicated 97.6% similarity to Myxobolus neurobius. Phylogenetic analysis revealed that M. compostellanus n. sp. is clustered with other salmonid-infecting myxobolids. The present findings contribute to the existing knowledge about the genus Myxobolus, providing both morphological and molecular data on a novel species of Myxobolus found to infect the gastrointestinal tract of salmonids, M. compostellanus n. sp. in the brown trout (S. trutta).

Negative associations between parasite avoidance, resistance and tolerance predict host health in salmonid fish populations.

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.

Modulation of local and systemic immune responses in brown trout (Salmo trutta) following exposure to Myxobolus cerebralis.

Myxobolus cerebralis, the etiological agent of Whirling Disease (WD), is a freshwater myxozoan parasite with considerable economic and ecological relevance for salmonids. There are differences in disease susceptibility between species and strains of salmonids. Recently, we have reported that the suppressor of cytokine signaling SOCS1 and SOCS3 are key in modulating rainbow trout (Oncorhynchus mykiss) immune responses and that resistant fish apparently exhibit effective Th17 cell response after exposure to M. cerebralis. It is unclear whether such molecules and pathways are also involved in the immune response of M. cerebralis infected brown trout (Salmo trutta). Hence, this study aimed to explore their role during immune modulation in infected brown trout, which is considered resistant to this parasite. Fish were exposed to the triactinomyxon (TAM) stages of M. cerebralis and quantitative real-time PCR (RT-qPCR) was carried out to examine local (caudal fin) and systemic (head kidney, spleen) immune transcriptional changes associated with WD over time in infected and control fish. All of the immune genes in the three tissues studied were differentially expressed in infected fish at multiple time points. Brown trout reduced the parasite load and demonstrated effective immune responses, likely by keeping pro-inflammatory and anti-inflammatory cytokines in balance whilst stimulating efficient Th17-mediated immunity. This study increases knowledge on the brown trout immune response to M. cerebralis and helps us to understand the underlying mechanisms of WD resistance.

Integrated field, laboratory, and theoretical study of PKD spread in a Swiss prealpine river.

Proliferative kidney disease (PKD) is a major threat to wild and farmed salmonid populations because of its lethal effect at high water temperatures. Its causative agent, the myxozoan Tetracapsuloides bryosalmonae, has a complex lifecycle exploiting freshwater bryozoans as primary hosts and salmonids as secondary hosts. We carried out an integrated study of PKD in a prealpine Swiss river (the Wigger). During a 3-year period, data on fish abundance, disease prevalence, concentration of primary hosts' DNA in environmental samples [environmental DNA (eDNA)], hydrological variables, and water temperatures gathered at various locations within the catchment were integrated into a newly developed metacommunity model, which includes ecological and epidemiological dynamics of fish and bryozoans, connectivity effects, and hydrothermal drivers. Infection dynamics were captured well by the epidemiological model, especially with regard to the spatial prevalence patterns. PKD prevalence in the sampled sites for both young-of-the-year (YOY) and adult brown trout attained 100% at the end of summer, while seasonal population decay was higher in YOY than in adults. We introduce a method based on decay distance of eDNA signal predicting local species' density, accounting for variation in environmental drivers (such as morphology and geology). The model provides a whole-network overview of the disease prevalence. In this study, we show how spatial and environmental characteristics of river networks can be used to study epidemiology and disease dynamics of waterborne diseases.

Field study indicating susceptibility differences between salmonid species and their lineages to proliferative kidney disease.

Tetracapsuloides bryosalmonae (Myxozoa: Malacosporea) is the causative agent of proliferative kidney disease (PKD), which affects both wild and farmed salmonid fish. The objective of this study was to outline differences in susceptibility to PKD in different salmonid species, hybrids and breeding lineages. Susceptibility to T. bryosalmonae infection was established based on cumulative mortality, pathological findings and detection of T. bryosalmonae in the kidney using immunohistochemistry and molecular methods. Determination of pure and hybrid individuals of different species in the genus Salvelinus, and dissimilarity of rainbow trout lineages, was performed using traditional polymerase chain reaction (PCR) and microsatellite analyses. Rainbow trout displayed higher disease severity compared with brook trout and Alsatian charr. Moreover, the results indicated differences in infection susceptibility, not only among different salmonid species but also among different lineages of charr and rainbow trout. Our study indicated that some salmonid species and even different lineages of the same species are more suitable for farming under PKD pressure.

Identification and Expression Profiling of Toll-Like Receptors of Brown Trout (Salmo trutta) during Proliferative Kidney Disease.

Proliferative kidney disease is an emerging disease among salmonids in Europe and North America caused by the myxozoan parasite Tetracapsuloides bryosalmonae. The decline of endemic brown trout (Salmo trutta) in the Alpine streams of Europe is fostered by T. bryosalmonae infection. Toll-like receptors (TLRs) are a family of pattern recognition receptors that acts as sentinels of the immune system against the invading pathogens. However, little is known about the TLRs' response in salmonids against the myxozoan infection. In the present study, we identified and evaluated TLR1, TLR19, and TLR13-like genes of brown trout using data-mining and phylogenetic analysis. The expression pattern of TLRs was examined in the posterior kidney of brown trout infected with T. bryosalmonae at various time points. Typical Toll/interleukin-1 receptor protein domain was found in all tested TLRs. However, TLR13-like chr2 had a short amino acid sequence with no LRR domain. Phylogenetic analysis illustrated that TLR orthologs are conserved across vertebrates. Similarly, a conserved synteny gene block arrangement was observed in the case of TLR1 and TLR19 across fish species. Interestingly, all tested TLRs showed their maximal relative expression from 6 to 10 weeks post-exposure to the parasite. Our results suggest that these TLRs may play an important role in the innate defense mechanism of brown trout against the invading T. bryosalmonae.

Saprolegnia molecular phylogeny among farmed teleosts in Nova Scotia, Canada.

To identify the pathogens causing saprolegniosis among farmed fish in Nova Scotia, 172 infected tissues and 23 water samples were collected from six species of teleosts: Atlantic salmon (Salmo salar), brown trout (Salmo trutta), Arctic charr (Salvelinus alpinus), brook trout (Salvelinus fontinalis), striped bass (Morone saxatilis) and rainbow trout (Oncorhynchus mykiss) at nine facilities over a 600 km range. Following laboratory culture, 132 isolates were recovered. Six species of oomycetes were identified from analysis of the internal transcribed spacer (ITS) sequence of the nrDNA: Saprolegnia parasitica, Saprolegnia ferax, Saprolegnia diclina, Saprolegnia aenigmatica, Saprolegnia torulosa, Saprolegnia sp. and Pythiopsis cymosa. Further phylogenetic analyses of the ITS and cytochrome c oxidase subunit 1 (Cox1) regions revealed four strains of Saprolegnia parasitica (named here as S1, S2, S3 and S4), of which S1 and S2 were common (37% and 42% of the isolates), and two strains of S. ferax. Among S. parasitica, S2 and S3 are more closely related to each other than to S1 based on the phylogenetic analyses and predicted RNA secondary structure of the ITS region. Sexual structures with a similar morphology were formed by S1 and S3 in vitro, but were not formed by S2.

Host (Salmo trutta) age influences resistance to infestation by freshwater pearl mussel (Margaritifera margaritifera) glochidia.

The freshwater pearl mussel (Margaritifera margaritifera) is an endangered bivalve with an obligate parasitic stage on salmonids. Host suitability studies have shown that glochidial growth and load vary significantly between host strains as well as among individuals of a suitable strain. Variation in host suitability has been linked to environmental conditions, host age and/or size, genetic composition of the host and parasite, or a combination of these factors. In our study, we wanted to investigate if brown trout (Salmo trutta) displayed an age-dependent response to glochidial infestation. We hypothesised that 1+ naive brown trout hosts tolerate glochidial infestation better than 0+ hosts. In order to test our hypothesis, we infested 0+ and 1+ hatchery reared brown trout with glochidia from closely related mothers and kept them under common garden conditions. This allowed us to observe a pure age dependent host response to infestation, as we eliminated the confounding effect of genotype-specific host interactions. We analysed the interaction between glochidial load and host condition, weight and length, and observed a significant age-dependent relationship. Glochidial load was negatively correlated to host condition in 0+ fish hosts and positively correlated in 1+ hosts. These contradictory findings can be explained by a change in host response strategy, from resistance in young to a higher tolerance in older fish. In addition, we also examined the relationship between glochidial load and haematocrit values in the 1+ hosts and observed that haematocrit values were significantly higher in heavily infested hosts. Our results have important conservation implications for the management of wild pearl mussel populations, as well as for captive breeding programmes.

First molecular data on Eimeria truttae from brown trout (Salmo trutta).

The genus Eimeria comprises obligate intracellular protozoan parasites belonging to the phylum Apicomplexa. Members of this genus cause enteric disease in a wide range of vertebrate hosts, including fish, reptiles, birds, and mammals. A total of 157 species of Eimeria that parasitize fish have been described; however, molecular information regarding these piscine parasites is scarce. In the present study, Eimeria oocysts were detected in 189 of 613 (30.8%) gastrointestinal tracts of brown trout (Salmo trutta) captured in several rivers in Galicia (NW Spain). Measurements of the sporulated oocysts, sporocysts, and other morphological characteristics enabled identification of the oocysts as Eimeria truttae. By molecular analysis of the small subunit ribosomal RNA (SSU-rRNA) gene, a single sequence of ~ 420 bp was obtained in 100 fish samples. After amplification of a ~ 1300-bp fragment of the same locus, two representative sequences that exhibited five nucleotide differences were obtained. Phylogenetic analysis grouped the samples within the piscine clade closest to Eimeria nemethi as they exhibited 96.7% similarity with this species. This study is the first to characterize E. truttae at the molecular level, thus helping to clarify the phylogenetic relationships between this and other Eimeria species isolated from fish and contributing further to the knowledge about this protozoan parasite.

A new species of Myxobolus (Myxozoa: Bivalvulida) infecting the medulla oblongata and nerve cord of brook trout Salvelinus fontinalis in southern Appalachia (New River, NC, USA).

Myxobolus neurofontinalis n. sp. infects the brain and medulla oblongata of brook trout (Salvelinus fontinalis [Mitchill, 1814]) in the New River, western NC. It is the first species of Myxobolus described from the brook trout and resembles another congener (Myxobolus arcticus Pugachev and Khokhlov, 1979) that infects nerve tissue of chars (Salvelinus spp.). The new species differs from M. arcticus and all congeners by myxospore dimensions and by having a mucous envelope and distinctive sutural markings. A phylogenetic analysis of the small subunit rDNA (18S) suggests that the new species shares a recent common ancestor with some isolates identified as M. arcticus and that the new species and its close relatives (except Myxobolus insidiosus Wyatt and Pratt, 1973) comprise a clade of salmonid nerve-infecting myxobolids. The phylogenetic analysis indicates that several isolates of "M. arcticus" (sensu lato) in GenBank are misidentified and distantly related to other isolates taken from the type host (Oncorhynchus nerka [Walbaum, 1792]) and from nearby the type locality (Kamchatka Peninsula, Russia). Serial histological sections of infected brook trout confirmed that myxospores of the new species are intercellular and infect nerve cord and medulla oblongata only. A single infected brook trout showed an inflammatory response characterized by focal lymphocytic infiltrates and eosinophilic granulocytes; however, the remaining 4 brook trout lacked evidence of a histopathological change or demonstrable host response. These results do not support the notion that this infection is pathogenic among brook trout.

Determinants of parasite distribution in Arctic charr populations: catchment structure versus dispersal potential.

Parasite distribution patterns in lotic catchments are driven by the combined influences of unidirectional water flow and the mobility of the most mobile host. However, the importance of such drivers in catchments dominated by lentic habitats are poorly understood. We examined parasite populations of Arctic charr Salvelinus alpinus from a series of linear-connected lakes in northern Norway to assess the generality of lotic-derived catchment-scale parasite assemblage patterns. Our results demonstrated that the abundance of most parasite taxa increased from the upper to lower catchment. Allogenic taxa (piscivorous birds as final host) were present throughout the entire catchment, whereas their autogenic counterparts (charr as final hosts) demonstrated restricted distributions, thus supporting the theory that the mobility of the most mobile host determines taxa-specific parasite distribution patterns. Overall, catchment-wide parasite abundance and distribution patterns in this lentic-dominated system were in accordance with those reported for lotic systems. Additionally, our study highlighted that upper catchment regions may be inadequate reservoirs to facilitate recolonization of parasite communities in the event of downstream environmental perturbations.

Natural selection for body shape in resource polymorphic Icelandic Arctic charr.

Resource polymorphisms exhibit remarkable intraspecific diversity and in many cases are expected to be maintained by diversifying selection. Phenotypic trade-offs can constrain morphologically intermediate individuals from effectively exploiting both alternate resources, resulting in ecological barriers to gene flow. Determining if and how phenotypic trade-offs cause fitness variation in the wild is challenging because of phenotypic and environmental correlations associated with alternative resource strategies. We investigated multiple pathways through which morphology could affect organismal performance, as measured by growth rate, and whether these effects generate diversifying selection in polymorphic Icelandic Arctic charr (Salvelinus alpinus) populations. We considered direct effects of morphology on growth and indirect effects via trophic resource use, estimated by stable isotopic signatures, and via parasitism associated with trophic resources. We sampled over 3 years in (lakes) Thingvallavatn and Vatnshlíðarvatn using the extended selection gradient path analytical approach and estimating size-dependent mortality. We found evidence for diversifying selection only in Thingvallavatn: more streamlined and terminally mouthed planktivore charr experienced greater growth, with the opposite pattern in small benthic charr. However, this effect was mediated by parasitism and nontrophic pathways, rather than trophic performance as often expected. Detection of between-morph differences in the presence (Vatnshlíðarvatn) and direction (Thingvallavatn) of size-dependent mortality, together with nontrophic effects of shape, suggests that a morphological trophic performance explanation for polymorphism is insufficient. This rare insight into selection during early diversification suggests that a complex of interacting local factors must be considered to understand how phenotype influences fitness, despite morphological variation reflecting intuitive trade-off explanations.

Temperature-related parasite infection dynamics: the case of proliferative kidney disease of brown trout.

Climate change, in particular rising temperature, is suspected to be a major driver for the emergence of many wildlife diseases. Proliferative kidney disease of salmonids, caused by the myxozoan Tetracapsuloides bryosalmonae, was used to evaluate how temperature dependence of host-parasite interactions modulates disease emergence. Brown trout (Salmo trutta fario) kept at 12 and 15 °C, were experimentally infected with T. bryosalmonae. Parasite development in the fish host and release of spores were quantified simultaneously to unravel parasite transmission potential from the vertebrate to the invertebrate host. A change to a stable plateau in infection intensity of the kidney coincided with a threshold at which spore shedding commenced. This onset of parasite release was delayed at the low temperature in accordance with reaching this infection intensity threshold, but the amount of spores released was irrespective of temperature. The production of parasite transmission stages declined with time. In conclusion, elevated temperature modifies the parasite transmission opportunities by increasing the duration of transmission stage production, which may affect the spread and establishment of the parasite in a wider range of rivers.