Consequences of reduced effectiveness of salmon lice treatments for lice control.

The effective control of ectoparasitic salmon lice, Lepeophtheirus salmonis, in fish farms is challenged by the salmon lice having developed resistance towards several antiparasitic drugs and by the effectiveness of non-medicinal treatments being limited by considerations of fish welfare. When new antiparasitics are introduced to the market, these should be used sparingly to slow resistance development. Using a population model for salmon lice parameterised for salmonid fish farms in Norway, we quantified how reduced treatment effectiveness influences treatment frequency and lice abundance. Furthermore, we investigated when in the production cycle a highly effective lice treatment leads to the largest reduction in the total number of treatments, mean lice abundance and lice larvae production. Results showed that reductions in treatment effectiveness to lower than 50% led to the steepest increases in treatment frequency and mean lice abundance, as well as to increased risk that lice abundance increased beyond control. The timing of the most effective treatment had only moderate effects on the total treatment need and the mean number of adult female lice through the production cycle, but large effect on the production of lice larvae in spring. These findings imply that farmers can optimise the timing of the most effective treatment to reduce the release of lice larvae in the period of year when wild salmonids are in coastal waters, without compromising total treatment need or mean lice levels.

Realtime case study simulations of transmission of Pancreas Disease (PD) in Norwegian salmonid farming for disease control purposes.

Pancreas Disease (PD) is a viral disease caused by Salmonid Alphavirus (SAV). It affects farmed salmonids in the North Atlantic, and leads to reduced feed intake and increased mortality with reduced production and welfare as a consequence. In 2013, the estimated cost of an outbreak on an average salmon farm was about 6.6 mil €. In Norway, PD has been notifiable since 2008, and regulations to mitigate disease spread are in place. However, despite the regulations, 140-170 farms are affected by PD every year. The aquaculture industry is growing continuously, introducing farms in new geographical areas, and fish are moved between hydrographically separated zones for trade and slaughter. All such movements and relocations need to be approved by the competent authorities. Thus, there is a demand for support to farmers and competent authorities when making decisions on disease management and especially on the effect of moving infected fish. We have used a disease-transmission model for outbreak-simulation in real time for assessing the probability of disease transmission from a farm that gets infected with PD. We have also simulated the effects of three different control-regimes: no stamping-out, delayed stamping-out or immediate stamping-out, on the transmission of PD to surrounding farms. Simulations showed that the immediate stamping out of an infected farm led to effective containment of an outbreak. No stamping out led to up to 32.1% of farms within 100 km of the index farm to become effected. We have used real production data for the model building and the scenario simulations, and the results illustrate that a risk assessment of horizontal disease transmission must be undertaken on a case-by-case basis, because the time and place of the outbreak has a large influence on the risk of transmission.

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.

Infectious Salmon Anemia Virus Shedding from Infected Atlantic Salmon (Salmo salar L.)-Application of a Droplet Digital PCR Assay for Virus Quantification in Seawater.

Infectious salmon anemia virus (ISAV) infection is currently detected by fish sampling for PCR and immunohistochemistry analysis. As an alternative to sampling fish, we evaluated two different membrane filters in combination with four buffers for elution, concentration, and detection of ISAV in seawater, during a bath challenge of Atlantic salmon (Salmo salar L.) post-smolts with high and low concentrations of ISAV. Transmission of ISAV in the bath challenge was confirmed by a high mortality, clinical signs associated with ISA disease, and detection of ISAV RNA in organ tissues and seawater samples. The electronegatively charged filter, combined with lysis buffer, gave significantly higher ISAV RNA detection by droplet digital PCR from seawater (5.6 × 104 ISAV RNA copies/L; p < 0.001). Viral shedding in seawater was first detected at two days post-challenge and peaked on day 11 post-challenge, one day before mortalities started in fish challenged with high dose ISAV, demonstrating that a large viral shedding event occurs before death. These data provide important information for ISAV shedding that is relevant for the development of improved surveillance tools based on water samples, transmission models, and management of ISA.

Early detection of salmonid alphavirus in seawater from marine farm sites of Atlantic salmon Salmo salar.

The traditional strategy for national surveillance of salmonid alphavirus (SAV) infection in Norwegian fish farms relies on a costly, time-consuming, and resource-demanding approach based on the monthly sampling of fish from all marine farms with salmonids. In order to develop an alternative surveillance method, a water filtration method was tested in parallel with the ongoing surveillance program at 7 Norwegian marine farm sites of Atlantic salmon Salmo salar L. with no current suspicion of SAV infection. During the period from May 2019 to January 2020, seawater samples were collected from the top layer water inside all net-pens at these 7 sites. The samples were concentrated for SAV by filtration through an MF-Millipore™ electronegative membrane filter, followed by rinsing with NucliSENS® Lysis Buffer, before RNA extraction and analysis by RT-qPCR. SAV was detected from seawater at an earlier stage compared to traditional sampling methods, at all sites where the fish tested positive for SAV. A significant negative relationship was observed at all sites between the SAV concentration found in seawater samples and the number of days until SAV was detected in the fish. This means that the fewer the SAV particles in the seawater, the more days it took until SAV was detected in the fish samples. Based on this, sampling of seawater every month for the surveillance of SAV has a great potential as an alternative method for early detection of SAV in Atlantic salmon farms.

Factors associated with baseline mortality in Norwegian Atlantic salmon farming.

In 2019, it was estimated that more than 50 million captive Atlantic salmon in Norway died in the final stage of their production in marine cages. This mortality represents a significant economic loss for producers and a need to improve welfare for farmed salmon. Single adverse events, such as algal blooms or infectious disease outbreaks, can explain mass mortality in salmon cages. However, little is known about the production, health, or environmental factors that contribute to their baseline mortality during the sea phase. Here we conducted a retrospective study including 1627 Atlantic salmon cohorts put to sea in 2014-2019. We found that sea lice treatments were associated with Atlantic salmon mortality. In particular, the trend towards non-medicinal sea lice treatments, including thermal delousing, increases Atlantic salmon mortality in the same month the treatment is applied. There were differences in mortality among production zones. Stocking month and weight were other important factors, with the lowest mortality in smaller salmon stocked in August-October. Sea surface temperature and salinity also influenced Atlantic salmon mortality. Knowledge of what affects baseline mortality in Norwegian aquaculture can be used as part of syndromic surveillance and to inform salmon producers on farming practices that can reduce mortality.

Filtration, concentration and detection of salmonid alphavirus in seawater during a post-smolt salmon (Salmo salar) cohabitant challenge.

Currently, the prevalence of salmonid alphavirus (SAV) in Norwegian Atlantic salmon farms is largely surveyed via sacrificing fish and sampling of organ tissue on a monthly basis. However, a more cost-efficient, straightforward, rapid, reliable, reproducible and animal welfare friendly method based on the detection of SAV in water could be considered as an alternative method. In the present study, such a method was developed and optimized through a 6 wk cohabitant challenge trial, using post-smolt Atlantic salmon Salmo salar L challenged with high or low doses of SAV subtype 3 (SAV3). Tank water and tissue samples from cohabitant fish were collected at 16 time points. SAV3 was concentrated from the water by filtration, using either electronegative or electropositive membrane filters, which were subsequently rinsed with one of 4 different buffer solutions. SAV3 was detected first in tank water (7 d post-challenge, DPC), and later in cohabitant fish organ tissue samples (12 DPC). The electronegative filter (MF-Millipore™) and rinsing with NucliSENS® easyMAG® Lysis Buffer presented the best SAV3 recovery. A significant positive correlation was found between SAV3 in the tank water concentrates and the mid-kidney samples. Based on these results, detection of SAV3 in filtrated seawater is believed to have the potential to serve as an alternative method for surveillance of SAV in Atlantic salmon farms.

Development and evaluation of a method for concentration and detection of salmonid alphavirus from seawater.

Waterborne viral infections represent a major threat to fish health. For many viruses, understanding the interplay between pathogens, host and environment presents a major hurdle for transmission. Salmonid alphavirus (SAV) can infect and cause pancreas disease (PD) in farmed salmonids in seawater. During infection, SAV is excreted from infected fish to the seawater. We evaluated two types of filters and four different eluents, for concentration of SAV3. One L of seawater was spiked with SAV3, followed by filtration and virus elution from membrane filters. For the negatively charged MF hydrophilic membrane filter (MF-) combined with NucliSENS® lysis buffer the SAV3 recovery was 39.5 ±â€¯1.8 % by RT-ddPCR and 25.9 ±â€¯5.7 % by RT-qPCR. The recovery using the positively charged 1 MDS Zeta Plus® Virosorb® membrane filter (MD+), combined with NucliSENS® lysis buffer was 19.0 ±â€¯0.1 % by RT-ddPCR and 13.3 ±â€¯3.8 % by RT-qPCR. The limits of quantification (LOQ) and detection (LOD) were estimated to be 5.18 × 103 and 2.0 × 102 SAV3 copies/L of natural seawater, by RT-ddPCR. SAV3 recovery from small volumes of seawater, and the requirement for standard laboratory equipment, suggest the MF-filter combined with NucliSENS® lysis buffer would be a candidate for further validation in experimental trials.

Spatio-temporal variations in mortality during the seawater production phase of Atlantic salmon (Salmo salar) in Norway.

In a sustainable production of animals, monitoring and minimizing mortality must be a top priority. Systematic measuring of mortality over time can be used to evaluate the impact of changes in management and production strategies in Norway. To aid understanding of the potential for reducing mortality, we have used data from 2014 to 2018 to investigate the spatio-temporal patterns of mortality, by descriptive analyses and statistical modelling of possible determinants of mortality. The results show large differences in mortality across different production zones and between years. The areas with the highest density of farmed salmon are also the ones with highest mortality. The total cumulated mortality of farmed salmon increased from 32.3 million in 2014 to 35.2 million in 2018, corresponding to 14.3% and 15.8% of the standing stock. An initial higher mortality was observed during the first 3 months after stocking (mean: 1.5% [0.9%-8.6%] mortality/month). This was followed by a period of stability and lower mortality (mean: 0.8% [0.9%-3.1%] mortality/month), until month 10, when mortality started to increase again. The month of first stocking, the year of slaughter, production zone and number of months at sea were all found to be statistically significant determinants for mortality, with p-values < 1e-15.

Infectious Salmon Anemia and Farm-Level Culling Strategies.

Infectious salmon anemia (ISA) is an infectious disease, and outbreaks must be handled to avoid spread between salmon sea farms. Intensive culling at infected farms is an important biosecurity measure to avoid further spread but is also a costly intervention that farmers try to avoid. A lack of action, however, may lead to new outbreaks in nearby salmon sea farms, with severe impacts on both economy and animal welfare. Here, we aim to explore how a time delay between a detected outbreak and the culling of both infected cages and entire farms affects the further spread of the disease. We use a previously published model to calculate how many salmon sea farms were directly infected in each outbreak. To investigate the effect of culling on the further spread of disease, we use the number of months elapsed from the detected outbreak to (a) the first cage being depopulated, and (b) to the entire salmon sea farm being depopulated as predictors of how many new farms the virus was transmitted to, after controlling for contact between the farms. We show that the lapse in time before the first cage is depopulated correlates positively with how many new salmon sea farms are infected, indicating that infected cages should be culled with as little time delay as possible. The model does not have sufficient power to separate between culling of only cages assumed to be infected and the entire farm, and, consequently, provides no direct empirical evidence for the latter. Lack of evidence is not evidence, however, and we argue that a high probability of spread between cages in infected salmon sea farms still supports the depopulation of entire farms as the safest option.

Risk Factors Associated With Outbreaks of Infectious Salmon Anemia (ISA) With Unknown Source of Infection in Norway.

The occurrence of infectious salmon anemia (ISA) outbreaks in marine farmed Atlantic salmon constitutes a recurring challenge in Norway. Here, we aim to identify risk factors associated with ISA outbreaks with an unknown source of infection (referred to as primary ISA outbreaks). Primary ISA outbreaks are here defined by an earlier published transmission model. We explored a wide range of possible risk factors with logistic regression analysis, trying to explain occurrence of primary ISA with available data from all Norwegian farm sites from 2004 to June 2017. Explanatory variables included site latitude and a range of production and disease data. The mean annual risk of having a primary outbreak of ISA in Norway was 0.7% during this study period. We identified the occurrence of infectious pancreatic necrosis (IPN), having a stocking period longer than 2 months, having the site located at high latitude and high fish density (biomass per cage volume) in the first six months after transfer to sea site as significant risk factors (p < 0.05). We have identified factors related to management routines, other disease problems, and latitude that may help to understand the hitherto unidentified drivers behind the emergence of primary ISA outbreaks. Based on our findings, we also provide management advice that may reduce the incidence of primary ISA outbreaks.

Parasitic sea louse infestations on wild sea trout: separating the roles of fish farms and temperature.

BACKGROUND: The causal relation between parasitic sea lice on fish farms and sea lice on wild fish is a controversial subject. A specific scientific debate has been whether the statistical association between infestation pressure (IP) from fish farms and the number of parasites observed on wild sea trout emerges purely because of a confounding and direct effect of temperature (T). METHODS: We studied the associations between louse infestation on wild sea trout, fish farm activity and temperature in an area that practices coordinated fallowing in Nordhordland, Norway. The data were sampled between 2009 and 2016. We used negative binomial models and mediation analysis to determine to what degree the effect of T is mediated through the IP from fish farms. RESULTS: The number of attached lice on sea trout increased with the T when the IP from fish farms was high but not when the IP was low. In addition, nearly all of the effect of rising T was indirect and mediated through the IP. Attached lice remained low when neighbouring farms were in the first year of the production cycle but rose substantially during the second year. In contrast to attached lice, mobile lice were generally seen in higher numbers at lower water temperatures. Temperature had an indirect positive effect on mobile louse counts by increasing the IP which, in turn, raised the sea trout louse counts. Mobile louse counts rose steadily during the year when neighbouring farms were in the first year of the production cycle and stayed high throughout the second year. CONCLUSIONS: The estimates of the IP effect on louse counts along with the clear biennial pattern emerging due to the production cycle of fish farms clearly indicate that fish farms play an important role in the epidemiology of sea lice on wild sea trout. Furthermore, the mediation analysis demonstrates that a large proportion of the effect of T on louse counts is mediated through IP.

Evaluation of aquaculture management zones as a control measure for salmon lice in Norway.

We evaluated the use of coordinated fallowing as a means to control salmon lice Lepeophtheirus salmonis infestation in farmed Atlantic salmon Salmo salar. In discrete management zones, aquaculture operations such as stocking, fallowing, treatments and harvesting are synchronized at all sites in coordinated areas within the zones. The expected benefit of synchronized generations is to reduce the presence of salmon lice larvae after a period of fallowing, as well as to minimize external infestation pressure from surrounding aquaculture sites. A regression analysis was used to evaluate the effectiveness of coordinated fallowing on the progression of external salmon lice infestation pressure and abundance in Atlantic salmon farming sites in 2 areas (zones) in Norway. The overall results show that external infestation pressure was higher inside than outside the management zones, and the external infestation pressure increased with increasing biomass throughout the production cycle. However, within the zones, the external infestation pressure at the beginning of a production cycle was high and in many cases even higher than the general external infestation pressure in the non-coordinated areas. This suggests that external infestation pressure from the neighboring areas has a considerable effect on the fallowed area. Higher numbers of salmon lice were recorded within the zones than outside and, as the production cycle progressed, this phenomenon became more evident. We conclude that the high infestation pressure from salmon lice at the beginning of the grow-out period after fallowing raises severe doubts about the effectiveness of coordinated fallowing practices.

Quantitative risk assessment of salmon louse-induced mortality of seaward-migrating post-smolt Atlantic salmon.

The Norwegian government recently implemented a new management system to regulate salmon farming in Norway, aiming to promote environmentally sustainable growth in the aquaculture industry. The Norwegian coast has been divided into 13 production zones and the volume of salmonid production in the zones will be regulated based on salmon lice effects on wild salmonids. Here we present a model for assessing salmon louse-induced mortality of seaward-migrating post-smolts of Atlantic salmon. The model quantifies expected salmon lice infestations and louse-induced mortality of migrating post-smolt salmon from 401 salmon rivers draining into Norwegian coastal waters. It is assumed that migrating post-smolts follow the shortest path from river outlets to the high seas, at constant progression rates. During this migration, fish are infested by salmon lice of farm origin according to an empirical infestation model. Furthermore, louse-induced mortality is estimated from the estimated louse infestations. Rivers draining into production zones on the West Coast of Norway were at the highest risk of adverse lice effects. In comparison, rivers draining into northerly production zones, along with the southernmost production zone, were at lower risk. After adjusting for standing stock biomass, estimates of louse-egg output varied by factors of up to 8 between production zones. Correlation between biomass adjusted output of louse infestation and densities of farmed salmon in the production zones suggests that a large-scale density-dependent host-parasite effect is a major driver of louse infestation rates and parasite-induced mortality. The estimates are sensitive to many of the processes in the chain of events in the model. Nevertheless, we argue that the model is suited to assess spatial and temporal risks associated with farm-origin salmon lice.

The influence of red deer space use on the distribution of Ixodes ricinus ticks in the landscape.

BACKGROUND: Many wingless ectoparasites have a limited capacity for active movement and are therefore primarily dependent on hitchhiking on their hosts for transportation. The distribution of the tick Ixodes ricinus is expected to depend mainly on transportation by hosts and tick subsequent survival in areas where they drop off. In Europe, the most important hosts of adult female I. ricinus are cervids. The extensive space use of large hosts provides a much larger dispersal potential for I. ricinus than that of smaller mammalian hosts. We aim to determine the contribution of red deer (Cervus elaphus) space use on the spatial distribution of I. ricinus, after accounting for landscape factors. METHODS: We analysed the spatial distribution of I. ricinus with generalised mixed effects models (GLMMs) based on data from extensive field surveys of questing density in two coastal regions in Norway, from which home range data from 73 red deer with GPS collars were available. Red deer home ranges were derived using the kernel method to identify areas most frequently used by deer. We first fitted a baseline model with tick questing densities relative to landscape features that are likely to affect local climate conditions and hence, survival. We then added deer space use variables to the baseline model with only landscape variables to test whether areas more frequently used by red deer had higher questing tick densities. RESULTS: Questing I. ricinus density was predicted by several landscape features, such as elevation, distance to the fjord and topographic slope. In addition, we found that areas more heavily used within the red deer home ranges, correlated with higher questing tick densities. Increased effects of deer space use were additive to the landscape model, suggesting that correlations were more than just shared landscape preferences between deer and ticks. CONCLUSIONS: Our results imply that the distribution of I. ricinus is controlled by a complex set of factors that include both local conditions related to landscape properties that affect survival and how the large host population redistributes ticks. In particular, we have provided evidence that the local distribution of large hosts, with their extensive space use, redistributes ticks at the local scale.

Parasite load and seasonal migration in red deer.

Northern deer populations are typically partially migratory, but the relationship between migratory movements and parasites has received little attention. Migration often involves movement from a low-elevation winter range towards a summer range at higher elevation. In Europe these movements may also involve a gradient in abundance of Ixodes ricinus ticks, but whether tick loads on deer differ depending on migration tactic has not been quantified. Based on the examination of ears from 49 red deer (Cervus elaphus) marked with global positioning system collars, we provide the first evidence that the tick loads of deer covering longer distances between their winter and summer range, resulting in higher difference in elevation, are lower. Our study highlights that only the resident part of the red deer population will be available as year-round hosts to ticks, while a large part of the red deer population is unavailable to ticks for most of the tick questing season due to seasonal migration to higher elevation. Predicted changes in the migratory behaviour of ungulates could hence affect the proportion of the host population available to ticks in the future.

The generalist tick Ixodes ricinus and the specialist tick Ixodes trianguliceps on shrews and rodents in a northern forest ecosystem--a role of body size even among small hosts.

BACKGROUND: Understanding aggregation of ticks on hosts and attachment of life stages to different host species, are central components for understanding tick-borne disease epidemiology. The generalist tick, Ixodes ricinus, is a well-known vector of Lyme borrelioses, while the specialist tick, Ixodes trianguliceps, feeding only on small mammals, may play a role in maintaining infection levels in hosts. In a northern forest in Norway, we aimed to quantify the role of different small mammal species in feeding ticks, to determine the extent to which body mass, even among small mammals, plays a role for tick load, and to determine the seasonal pattern of the two tick species. METHODS: Small mammals were captured along transects in two nearby areas along the west coast of Norway. All life stages of ticks were counted. Tick load, including both prevalence and intensity, was analysed with negative binomial models. RESULTS: A total of 359 rodents and shrews were captured with a total of 1106 I. ricinus (60.0 %) and 737 I. trianguliceps (40.4 %), consisting of 98.2 % larvae and 1.8 % nymphs of I. ricinus and 91.2 % larvae, 8.7 % nymphs and 0.1 % adult females of I. trianguliceps. Due to high abundance, Sorex araneus fed most of the larvae of both tick species (I. ricinus 61.9 %, I. trianguliceps 64.9 %) with Apodemus sylvaticus (I. ricinus 20.4 %, I. trianguliceps 10.0 %) and Myodes glareolus (I. ricinus 10.9 %, I. trianguliceps 9.5 %) as the next most important hosts. Individual A. sylvaticus and M. glareolus had higher infestation intensity than S. araneus, while Sorex minutus had markedly lower infestation intensity. The load of I. ricinus larvae and nymphs was related to body mass mainly up to ~10 g, while the load of I. trianguliceps was less dependent of body mass. The load of I. trianguliceps was higher in spring than in fall, while the seasonal pattern was reversed for I. ricinus with higher loads in fall. CONCLUSIONS: Body mass was important for explaining load of I. ricinus mainly up to a body mass of ~10 g across a range of smaller mammalian hosts. Consistent with earlier work elsewhere in Europe, we found the highest tick infestation intensity on the wood mouse A. sylvaticus. However, this rodent species fed only 20.4 % of all I. ricinus larvae, while the much more abundant S. araneus fed 61.9 %. Our study emphasizes an important quantitative role of the common shrew S. araneus as a main host to I. ricinus larvae and to both I. trianguliceps larvae and nymphs. The partly seasonal distinct attachment pattern of I. ricinus and I. trianguliceps is evidence for niche separation.

Temporal pattern of questing tick Ixodes ricinus density at differing elevations in the coastal region of western Norway.

BACKGROUND: Climate change can affect the activity and distribution of species, including pathogens and parasites. The densities and distribution range of the sheep tick (Ixodes ricinus) and it's transmitted pathogens appears to be increasing. Thus, a better understanding of questing tick densities in relation to climate and weather conditions is urgently needed. The aim of this study was to test predictions regarding the temporal pattern of questing tick densities at two different elevations in Norway. We predict that questing tick densities will decrease with increasing elevations and increase with increasing temperatures, but predict that humidity levels will rarely affect ticks in this northern, coastal climate with high humidity. METHODS: We described the temporal pattern of questing tick densities at ~100 and ~400 m a.s.l. along twelve transects in the coastal region of Norway. We used the cloth lure method at 14-day intervals during the snow-free season to count ticks in two consecutive years in 20 m2 plots. We linked the temporal pattern of questing tick densities to local measurements of the prevailing weather. RESULTS: The questing tick densities were much higher and the season was longer at ~100 compared to at ~400 m a.s.l. There was a prominent spring peak in both years and a smaller autumn peak in one year at ~100 m a.s.l.; but no marked peak at ~400 m a.s.l. Tick densities correlated positively with temperature, from low densities <5°C, then increasing and levelling off >15-17°C. We found no evidence for reduced questing densities during the driest conditions measured. CONCLUSIONS: Tick questing densities differed even locally linked to elevation (on the same hillside, a few kilometers apart). The tick densities were strongly hampered by low temperatures that limited the duration of the questing seasons, whereas the humidity appeared not to be a limiting factor under the humid conditions at our study site. We expect rising global temperatures to increase tick densities and lead to a transition from a short questing season with low densities in the current cold and sub-optimal tick habitats, to longer questing seasons with overall higher densities and a marked spring peak.

Landscape level variation in tick abundance relative to seasonal migration in red deer.

Partial migration is common among northern ungulates, typically involving an altitudinal movement for seasonally migratory individuals. The main driving force behind migration is the benefit of an extended period of access to newly emerged, high quality forage along the green up gradient with increasing altitude; termed the forage maturation hypothesis. Any other limiting factor spatially correlated with this gradient may provide extra benefits or costs to migration, without necessarily being the cause of it. A common ectoparasite on cervids in Europe is the sheep tick (Ixodes ricinus), but it has not been tested whether migration may lead to the spatial separation from these parasites and thus potentially provide an additional benefit to migration. Further, if there is questing of ticks in winter ranges in May before spring migration, deer migration may also play a role for the distribution of ticks. We quantified the abundance of questing sheep tick within winter and summer home ranges of migratory (n=42) and resident red deer (Cervus elaphus) individuals (n=32) in two populations in May and August 2009-2012. Consistent with predictions, there was markedly lower abundance of questing ticks in the summer areas of migrating red deer (0.6/20 m(2)), both when compared to the annual home range of resident deer (4.9/20 m(2)) and the winter home ranges of migrants (5.8/20 m(2)). The reduced abundances within summer home ranges of migrants were explained by lower abundance of ticks with increasing altitude and distance from the coast. The lower abundance of ticks in summer home ranges of migratory deer does not imply that ticks are the main driver of migration (being most likely the benefits expected from forage maturation), but it suggests that ticks may add to the value of migration in some ecosystems and that it may act to spread ticks long distances in the landscape.

Spatial and seasonal variation in the prevalence of Anaplasma phagocytophilum and Borrelia burgdorferi sensu lato in questing Ixodes ricinus ticks in Norway.

BACKGROUND: Understanding the variation in prevalence of Borrelia burgdorferi sensu lato (Lyme Borreliosis Spirochaetes, LBS) and Anaplasma phagocytophilum (causing tick-borne fever in ruminants and human granulocytic ehrlichiosis) in ticks is vital from both a human and an animal disease perspective to target the most effective mitigation measures. From the host competence hypothesis, we predicted that prevalence of LBS would decrease with red deer density, while prevalence of A. phagocytophilum would increase. METHODS: Based on a sample of 112 adult and 686 nymphal Ixodes ricinus ticks collected with flagging during questing from 31 transects (4-500 m long) corresponding to individual seasonal home ranges of 41 red deer along the west coast of Norway, we tested whether there were spatial and seasonal variations in prevalence with a special emphasis on the population density of the most common large host in this area, the red deer (Cervus elaphus). We used a multiplex real-time PCR assay for detection of A. phagocytophilum and LBS. RESULTS: Prevalence of LBS was higher in adult female ticks (21.6%) compared to adult male ticks (11.5%) and nymphs (10.9%), while prevalence was similar among stages for prevalence of A. phagocytophilum (8.8%). Only partly consistent with predictions, we found a lower prevalence of LBS in areas of high red deer density, while there was no relationship between red deer density and prevalence of A. phagocytophilum in ticks. Prevalence of both bacteria was much higher in ticks questing in May compared to August. CONCLUSIONS: Our study provides support to the notion that spatial variation in host composition forms a role for prevalence of LBS in ticks also in a northern European ecosystem, while no such association was found for A. phagocytophilum. Further studies are needed to fully understand the similar seasonal pattern of prevalence of the two pathogens.