Bottom-up rather than top-down mechanisms determine mesocarnivore interactions in Norway.

Interactions among coexisting mesocarnivores can be influenced by different factors such as the presence of large carnivores, land-use, environmental productivity, or human disturbance. Disentangling the relative importance of bottom-up and top-down processes can be challenging, but it is important for biodiversity conservation and wildlife management. The aim of this study was to assess how the interactions among mesocarnivores (red fox Vulpes vulpes, badger Meles meles, and pine marten Martes martes) were affected by large carnivores (Eurasian lynx Lynx lynx and wolf Canis lupus), land cover variables (proportion of agricultural land and primary productivity), and human disturbance, as well as how these top-down and bottom-up mechanisms were influenced by season. We analyzed 3 years (2018-2020) of camera trapping observations from Norway and used structural equation models to assess hypothesized networks of causal relationships. Our results showed that land cover variables were more strongly associated with mesocarnivore detection rates than large carnivores in Norway. This might be caused by a combination of low density of large carnivores in an unproductive ecosystem with strong seasonality. Additionally, detection rates of all mesocarnivores showed positive associations among each other, which were stronger in winter. The prevalence of positive interactions among predators might indicate a tendency to use the same areas and resources combined with weak interference competition. Alternatively, it might indicate some kind of facilitative relationship among species. Human disturbance had contrasting effects for different species, benefiting the larger mesocarnivores (red fox and badger) probably through food subsidization, but negatively affecting apex predators (wolf and lynx) and smaller mesocarnivores (pine marten). In a human-dominated world, this highlights the importance of including anthropogenic influences in the study of species interactions.

Snow cover-related camouflage mismatch increases detection by predators.

Camouflage expressed by animals is an adaptation to local environments that certain animals express to maximize survival and fitness. Animals at higher latitudes change their coat color according to a seasonally changing environment, expressing a white coat in winter and a darker coat in summer. The timing of molting is tightly linked to the appearance and disappearance of snow and is mainly regulated by photoperiod. However, due to climate change, an increasing mismatch is observed between the coat color of these species and their environment. Here, we conducted an experiment in northern Sweden, with white and brown decoys to study how camouflage (mis)-match influenced (1) predator attraction to decoys, and (2) predation events. Using camera trap data, we showed that mismatching decoys attracted more predators and experienced a higher likelihood of predation events in comparison to matching decoys, suggesting that camouflage mismatched animals experience increased detection by predators. These results provide insight into the function of a seasonal color coat and the need for this adaptation to maximize fitness in an environment that is exposed to high seasonality. Thus, our results suggest that, with increasing climate change and reduced snow cover, animals expressing a seasonal color coat will experience a decrease in survival.

Exploring the influence of host community composition on the outbreak potential of Anaplasma phagocytophilum and Borrelia burgdorferi s.l.

In large parts of the northern hemisphere, multiple deer species coexist, and management actions can strongly influence wild deer communities. Such changes may also indirectly influence other species in the community, such as small mammals and birds, because deer can have strong effects on their habitats and resources. Deer, small mammals and birds play an important role in the dynamics of tick-borne zoonotic diseases. It is, however, relatively underexplored how the abundance and composition of vertebrate communities may affect the outbreak potential, maintenance and circulation of tick-borne pathogens. In this study we focus on the outbreak potential by exploring how the basic reproduction number R0 for different tick-borne pathogens depends on host community composition. We used published data on co-varying roe deer (Capreolus capreolus) and fallow deer (Dama dama) densities following a hunting ban, and different small mammal and bird densities, to investigate how the change in host community influences the R0 of four tick-borne pathogens: one non-zoonotic, namely Anaplasma phagocytophilum ecotype 2, and three zoonotic, namely A. phagocytophilum ecotype 1, Borrelia afzelii and Borrelia garinii. We calculated R0 using a next generation matrix approach, and used elasticities to quantify the contributions to R0 of the different groups of host species. The value of R0 for A. phagocytophilum ecotype 1 was higher with high fallow deer density and low roe deer density, while it was the other way round for A. phagocytophilum ecotype 2. For B. afzelii, R0 was mostly related to the density of small mammals and for B. garinii it was mostly determined by bird density. Our results show that the effect of species composition is substantial in the outbreak potential of tick-borne pathogens. This implies that also management actions that change this composition, can (indirectly and unintentionally) affect the outbreak potential of tick-borne diseases.

Altitude, latitude and climate zone as determinants of mountain hare (Lepus timidus) coat colour change.

Local adaptation to annually changing environments has evolved in numerous species. Seasonal coat colour change is an adaptation that has evolved in multiple mammal and bird species occupying areas that experience seasonal snow cover. It has a critical impact on fitness as predation risk may increase when an individual is mismatched against its habitat's background colour. In this paper, we investigate the correlation between landscape covariates and moult timing in a native winter-adapted herbivore, the mountain hare (Lepus timidus), throughout Norway. Data was collected between 2011 and 2019 at 678 camera trap locations deployed across an environmental gradient. Based on this data, we created a Bayesian multinomial logistic regression model that quantified the correlations between landscape covariates and coat colour phenology and analysed among season and year moult timing variation. Our results demonstrate that mountain hare moult timing is strongly correlated with altitude and latitude with hares that live at higher latitudes and altitudes keeping their winter white coats for longer than their conspecifics that inhabit lower latitudes and altitudes. Moult timing was also weakly correlated with climate zone with hares that live in coastal climates keeping their winter white coats for longer than hares that live in continental climates. We found evidence of some among year moult timing variation in spring, but not in autumn. We conclude that mountain hare moult timing has adapted to local environmental conditions throughout Norway.

Smartphone app reveals that lynx avoid human recreationists on local scale, but not home range scale.

Outdoor recreation is increasing and affects habitat use and selection by wildlife. These effects are challenging to study, especially for elusive species with large spatial requirements, as it is hard to obtain reliable proxies of recreational intensity over extensive areas. Commonly used proxies, such as the density of, or distance to, hiking paths, ignore outdoor recreation occurring on other linear feature types. Here we utilized crowdsourced data from the Strava training app to obtain a large-scale proxy for pedestrian outdoor recreation intensity in southeast Norway. We used the proxy and GPS-tracking data from collared Eurasian lynx (Lynx lynx) to investigate how recreation affects habitat selection at the home range scale and local scale by lynx during summer. We fitted resource selection functions at the two scales using conditional logistic regression. Our analysis revealed that lynx avoided areas of recreational activity at the local scale, but not at home range scale. Nonetheless, lynx frequently used areas associated with recreation, and to a greater degree at night than during the day. Our results suggest that local-scale avoidance of recreation and temporal adjustments of habitat use by lynx mitigate the need for a home range-scale response towards recreation. Scale-dependent responses and temporal adjustments in habitat use may facilitate coexistence between humans and large carnivores.

Different increase rate in body mass of two marten species due to climate warming potentially reinforces interspecific competition.

Many species show spatial variation in body size, often associated with climatic patterns. Studying species with contrasting geographical patterns related to climate might help elucidate the role of different drivers. We analysed changes in the body mass of two sympatric medium-sized carnivores-pine marten (Martes martes) and stone marten (Martes foina)-across Europe over 59 years. The body mass of pine marten increased with decreasing latitude, whereas stone marten body mass varied in a more complex pattern across its geographic range. Over time, the average body mass of pine martens increased by 255 g (24%), while stone marten by 86 g (6%). The greatest increase of body mass along both martens' geographic range was observed in central and southern Europe, where both species occur in sympatry. The body mass increase slowed down over time, especially in allopatric regions. The average pine/stone marten body mass ratio increased from 0.87 in 1960 to 0.99 in 2019, potentially strengthening the competition between them. Thus, a differential response in body size to several drivers over time might have led to an adaptive advantage for pine martens. This highlights the importance of considering different responses among interacting species when studying animal adaptation to climate change.

Camera trap records of leucistic Eurasian badgers (Meles meles) in central Norway.

Coat coloration plays an important role in communication, camouflage, and sexual selection in animals. Genetic mutations can lead to anomalous colorations such as melanism and leucism, where animals appear, respectively, darker or lighter than normal. Reporting abnormal coloration in wild animals is an important first step to understand the distribution, prevalence, and potential fitness consequences of these rare events. Here, we report several records of suspected leucism in the Eurasian badger (Meles meles) in a population in central Norway. Several camera traps recorded at least two leucistic individuals between 2017 and 2020. It took considerable effort, almost 400,000 camera trap nights over a period of 10 years all over Norway, to obtain a total of eleven records of leucistic badgers, indicating the rarity of this phenotype. It is unclear what has caused the presence of multiple leucistic badgers in a single population, but recent colonization and lack of predators might have played a role. Due to our observations, future studies can now be developed to study the underlying mechanisms and potential consequences of leucism in this badger population. The increasing use of networks of camera traps in wildlife research will provide new opportunities to record rare coloration in wild animals.

Wild ungulate species differ in their contribution to the transmission of Ixodes ricinus-borne pathogens.

BACKGROUND: Several ungulate species are feeding and propagation hosts for the tick Ixodes ricinus as well as hosts to a wide range of zoonotic pathogens. Here, we focus on Anaplasma phagocytophilum and Borrelia burgdorferi (s.l.), two important pathogens for which ungulates are amplifying and dilution hosts, respectively. Ungulate management is one of the main tools to mitigate human health risks associated with these tick-borne pathogens. Across Europe, different species of ungulates are expanding their ranges and increasing in numbers. It is currently unclear if and how the relative contribution to the life-cycle of I. ricinus and the transmission cycles of tick-borne pathogens differ among these species. In this study, we aimed to identify these relative contributions for five European ungulate species. METHODS: We quantified the tick load and collected ticks and spleen samples from hunted fallow deer (Dama dama, n = 131), moose (Alces alces, n = 15), red deer (Cervus elaphus, n = 61), roe deer (Capreolus capreolus, n = 30) and wild boar (Sus scrofa, n = 87) in south-central Sweden. We investigated the presence of tick-borne pathogens in ticks and spleen samples using real-time PCR. We determined if ungulate species differed in tick load (prevalence and intensity) and in infection prevalence in their tissue as well as in the ticks feeding on them. RESULTS: Wild boar hosted fewer adult female ticks than any of the deer species, indicating that deer are more important as propagation hosts. Among the deer species, moose had the lowest number of female ticks, while there was no difference among the other deer species. Given the low number of infected nymphs, the relative contribution of all ungulate species to the transmission of B. burgdorferi (s.l.) was low. Fallow deer, red deer and roe deer contributed more to the transmission of A. phagocytophilum than wild boar. CONCLUSIONS: The ungulate species clearly differed in their role as a propagation host and in the transmission of B. burgdorferi and A. phagocytophilum. This study provides crucial information for ungulate management as a tool to mitigate zoonotic disease risk and argues for adapting management approaches to the local ungulate species composition and the pathogen(s) of concern.

Red and fallow deer determine the density of Ixodes ricinus nymphs containing Anaplasma phagocytophilum.

BACKGROUND: The density of Ixodes ricinus nymphs infected with Anaplasma phagocytophilum is one of the parameters that determines the risk for humans and domesticated animals to contract anaplasmosis. For this, I. ricinus larvae need to take a bloodmeal from free-ranging ungulates, which are competent hosts for A. phagocytophilum. METHODS: Here, we compared the contribution of four free-ranging ungulate species, red deer (Cervus elaphus), fallow deer (Dama dama), roe deer (Capreolus capreolus), and wild boar (Sus scrofa), to A. phagocytophilum infections in nymphs. We used a combination of camera and live trapping to quantify the relative availability of vertebrate hosts to questing ticks in 19 Dutch forest sites. Additionally, we collected questing I. ricinus nymphs and tested these for the presence of A. phagocytophilum. Furthermore, we explored two potential mechanisms that could explain differences between species: (i) differences in larval burden, which we based on data from published studies, and (ii) differences in associations with other, non-competent hosts. RESULTS: Principal component analysis indicated that the density of A. phagocytophilum-infected nymphs (DIN) was higher in forest sites with high availability of red and fallow deer, and to a lesser degree roe deer. Initial results suggest that these differences are not a result of differences in larval burden, but rather differences in associations with other species or other ecological factors. CONCLUSIONS: These results indicate that the risk for contracting anaplasmosis in The Netherlands is likely highest in the few areas where red and fallow deer are present. Future studies are needed to explore the mechanisms behind this association.

Predation risk constrains herbivores' adaptive capacity to warming.

Global warming compels larger endothermic animals to adapt either physiologically or behaviourally to avoid thermal stress, especially in tropical ecosystems. Their adaptive responses may however be compromised by other constraints, such as predation risk or starvation. Using an exceptional camera-trap dataset spanning 32 protected areas across southern Africa, we find that intermediate-sized herbivores (100-550 kg) switch activity to hotter times of the day when exposed to predation by lions. These herbivores face a tight window for foraging activity being exposed to nocturnal predation and to heat during the day, suggesting a trade-off between predation risk and thermoregulation mediated by body size. These findings stress the importance of incorporating trophic interactions into climate change predictions.

Effects of cattle grazing on Ixodes ricinus-borne disease risk in forest areas of the Netherlands.

Cattle grazing has been suggested to reduce the risk for Lyme borreliosis by decreasing the density of questing Ixodes ricinus infected with Borrelia burgdorferi sensu lato. We tested the hypotheses that cattle grazing used in woodland management decreases the density of questing I. ricinus, and that it decreases the nympal infection prevalence of B. burgdorferi sensu lato. We further expected the nympal infection prevalence of tick-borne pathogens that utilize cattle as amplifying hosts, namely Anaplasma phagocytophilum, and Babesia sensu stricto, to increase. To test these hypotheses, we compared the densities of questing I. ricinus between twenty pairs of plots in grazed and ungrazed forest areas. The density of I. ricinus adults, but not nymphs, was lower in areas grazed by cattle than in ungrazed areas. Nymphs were tested for the presence of Borrelia burgdorferi s.l., Borrelia miyamotoi, Anaplasma phagocytophilum, and Babesia s.s. DNA from twelve paired areas. Anaplasma phagocytophilum and Babesia s.s. from qPCR-positive tick lysates were identified further to the ecotype and species level, respectively, by DNA sequencing. The infection prevalence of A. phagocytophilum was lower, and infection prevalence of Babesia s.s., identified as Babesia venatorum, was higher in grazed areas. In contrast, infection prevalence with B. burgdorferi s.l. or B. miyamotoi did not differ between grazed and ungrazed areas. As a consequence, conventional cattle grazing in forested areas had no effect on the densities of questing nymphs infected with B. burgdorferi s.l. and B. miyamotoi. Similarly, we found no effect of cattle grazing on the density of infected nymphs with B. venatorum. The marked difference in the densities of questing nymphs infected with A. phagocytophilum could potentially be explained by the presence of a higher density of roe deer (Capreolus capreolus) in ungrazed areas, as the majority of typed A. phagocytophilum from ungrazed areas were the non-zoonotic ecotype II, which is associated with roe deer.

Parasite Load and Site-Specific Parasite Pressure as Determinants of Immune Indices in Two Sympatric Rodent Species.

Wildlife is exposed to parasites from the environment. This parasite pressure, which differs among areas, likely shapes the immunological strategies of animals. Individuals differ in the number of parasites they encounter and host, and this parasite load also influences the immune system. The relative impact of parasite pressure vs. parasite load on different host species, particularly those implicated as important reservoirs of zoonotic pathogens, is poorly understood. We captured bank voles (Myodes glareolus) and wood mice (Apodemus sylvaticus) at four sites in the Netherlands. We sampled sub-adult males to quantify their immune function, infestation load for ecto- and gastrointestinal parasites, and infection status for vector-borne microparasites. We then used regression trees to test if variation in immune indices could be explained by among-site differences (parasite pressure), among-individual differences in infestation intensity and infection status (parasite load), or other intrinsic factors. Regression trees revealed splits among sites for haptoglobin, hemagglutination, and body-mass corrected spleen size. We also found splits based on infection/infestation for haptoglobin, hemolysis, and neutrophil to lymphocyte ratio. Furthermore, we found a split between species for hemolysis and splits based on body mass for haptoglobin, hemagglutination, hematocrit, and body-mass corrected spleen size. Our results suggest that both parasite pressure and parasite load influence the immune system of wild rodents. Additional studies linking disease ecology and ecological immunology are needed to understand better the complexities of host-parasite interactions and how these interactions shape zoonotic disease risk.

Impact of vertebrate communities on Ixodes ricinus-borne disease risk in forest areas.

BACKGROUND: The density of questing ticks infected with tick-borne pathogens is an important parameter that determines tick-borne disease risk. An important factor determining this density is the availability of different wildlife species as hosts for ticks and their pathogens. Here, we investigated how wildlife communities contribute to tick-borne disease risk. The density of Ixodes ricinus nymphs infected with Borrelia burgdorferi (sensu lato), Borrelia miyamotoi, Neoehrlichia mikurensis and Anaplasma phagocytophilum among 19 forest sites were correlated to the encounter probability of different vertebrate hosts, determined by encounter rates as measured by (camera) trapping and mathematical modeling. RESULT: We found that the density of any tick life stage was proportional to the encounter probability of ungulates. Moreover, the density of nymphs decreased with the encounter probability of hare, rabbit and red fox. The density of nymphs infected with the transovarially-transmitted B. miyamotoi increased with the density of questing nymphs and the encounter probability of bank vole. The density of nymphs infected with all other pathogens increased with the encounter probability of competent hosts: bank vole for Borrelia afzelii and N. mikurensis, ungulates for A. phagocytophilum and blackbird for Borrelia garinii and Borrelia valaisiana. The negative relationship we found was a decrease in the density of nymphs infected with B. garinii and B. valaisiana with the encounter probability of wood mouse. CONCLUSIONS: Only a few animal species drive the densities of infected nymphs in forested areas. There, foxes and leporids have negative effects on tick abundance, and consequently on the density of infected nymphs. The abundance of competent hosts generally drives the abundances of their tick-borne pathogen. A dilution effect was only observed for bird-associated Lyme spirochetes.

Framing pictures: A conceptual framework to identify and correct for biases in detection probability of camera traps enabling multi-species comparison.

Obtaining reliable species observations is of great importance in animal ecology and wildlife conservation. An increasing number of studies use camera traps (CTs) to study wildlife communities, and an increasing effort is made to make better use and reuse of the large amounts of data that are produced. It is in these circumstances that it becomes paramount to correct for the species- and study-specific variation in imperfect detection within CTs. We reviewed the literature and used our own experience to compile a list of factors that affect CT detection of animals. We did this within a conceptual framework of six distinct scales separating out the influences of (a) animal characteristics, (b) CT specifications, (c) CT set-up protocols, and (d) environmental variables. We identified 40 factors that can potentially influence the detection of animals by CTs at these six scales. Many of these factors were related to only a few overarching parameters. Most of the animal characteristics scale with body mass and diet type, and most environmental characteristics differ with season or latitude such that remote sensing products like NDVI could be used as a proxy index to capture this variation. Factors that influence detection at the microsite and camera scales are probably the most important in determining CT detection of animals. The type of study and specific research question will determine which factors should be corrected. Corrections can be done by directly adjusting the CT metric of interest or by using covariates in a statistical framework. Our conceptual framework can be used to design better CT studies and help when analyzing CT data. Furthermore, it provides an overview of which factors should be reported in CT studies to make them repeatable, comparable, and their data reusable. This should greatly improve the possibilities for global scale analyses of (reused) CT data.

Role of mustelids in the life-cycle of ixodid ticks and transmission cycles of four tick-borne pathogens.

BACKGROUND: Elucidating which wildlife species significantly contribute to the maintenance of Ixodes ricinus populations and the enzootic cycles of the pathogens they transmit is imperative in understanding the driving forces behind the emergence of tick-borne diseases. Here, we aimed to quantify the relative contribution of four mustelid species in the life-cycles of I. ricinus and Borrelia burgdorferi (sensu lato) in forested areas and to investigate their role in the transmission of other tick-borne pathogens. Road-killed badgers, pine martens, stone martens and polecats were collected in Belgium and the Netherlands. Their organs and feeding ticks were tested for the presence of tick-borne pathogens. RESULTS: Ixodes hexagonus and I. ricinus were found on half of the screened animals (n = 637). Pine martens had the highest I. ricinus burden, whereas polecats had the highest I. hexagonus burden. We detected DNA from B. burgdorferi (s.l.) and Anaplasma phagocytophilum in organs of all four mustelid species (n = 789), and Neoehrlichia mikurensis DNA was detected in all species, except badgers. DNA from B. miyamotoi was not detected in any of the investigated mustelids. From the 15 larvae of I. ricinus feeding on pine martens (n = 44), only one was positive for B. miyamotoi DNA, and all tested negative for B. burgdorferi (s.l.), N. mikurensis and A. phagocytophilum. The two feeding larvae from the investigated polecats (n = 364) and stone martens (n = 39) were negative for all four pathogens. The infection rate of N. mikurensis was higher in feeding nymphs collected from mustelids compared to questing nymphs, but not for B. burgdorferi (s.l.), B. miyamotoi or A. phagocytophilum. CONCLUSIONS: Although all stages of I. ricinus can be found on badgers, polecats, pine and stone martens, their relative contribution to the life-cycle of I. ricinus in forested areas is less than 1%. Consequently, the relative contribution of mustelids to the enzootic cycles of I. ricinus-borne pathogens is negligible, despite the presence of these pathogens in organs and feeding ticks. Interestingly, all four mustelid species carried all stages of I. hexagonus, potentially maintaining enzootic cycles of this tick species apart from the cycle involving hedgehogs as main host species.

The genetic diversity of Borrelia afzelii is not maintained by the diversity of the rodent hosts.

BACKGROUND: Small mammals are essential in the enzootic cycle of many tick-borne pathogens (TBP). To understand their contribution to the genetic diversity of Borrelia afzelii, the most prevalent TBP in questing Ixodes ricinus, we compared the genetic variants of B. afzelii at three distinct genetic loci. We chose two plasmid loci, dbpA and ospC, and a chromosomal one, IGS. RESULTS: While the larvae that fed on shrews (Sorex sp.) tested negative for B. afzelii, those fed on bank voles (Myodes glareolus) and wood mice (Apodemus sylvaticus) showed high infection prevalences of 0.13 and 0.27, respectively. Despite the high genetic diversity within B. afzelii, there was no difference between wood mice and bank voles in the number and types of B. afzelii haplotypes they transmit. CONCLUSIONS: The genetic diversity in B. afzelii cannot be explained by separate enzootic cycles in wood mice and bank voles.

Deer presence rather than abundance determines the population density of the sheep tick, Ixodes ricinus, in Dutch forests.

BACKGROUND: Understanding which factors drive population densities of disease vectors is an important step in assessing disease risk. We tested the hypothesis that the density of ticks from the Ixodes ricinus complex, which are important vectors for tick-borne diseases, is determined by the density of deer, as adults of these ticks mainly feed on deer. METHODS: We performed a cross-sectional study to investigate I. ricinus density across 20 forest plots in the Netherlands that ranged widely in deer availability to ticks, and performed a deer-exclosure experiment in four pairs of 1 ha forest plots in a separate site. RESULTS: Ixodes ricinus from all stages were more abundant in plots with deer (n = 17) than in plots without deer (n = 3). Where deer were present, the density of ticks did not increase with the abundance of deer. Experimental exclosure of deer reduced nymph density by 66% and adult density by 32% within a timeframe of two years. CONCLUSIONS: In this study, deer presence rather than abundance explained the density of I. ricinus. This is in contrast to previous studies and might be related to the relatively high host-species richness in Dutch forests. This means that reduction of the risk of acquiring a tick bite would require the complete elimination of deer in species rich forests. The fact that small exclosures (< 1 ha) substantially reduced I. ricinus densities suggests that fencing can be used to reduce tick-borne disease risk in areas with high recreational pressure.

Quantifying the Availability of Vertebrate Hosts to Ticks: A Camera-Trapping Approach.

The availability of vertebrate hosts is a major determinant of the occurrence of ticks and tick-borne zoonoses in natural and anthropogenic ecosystems and thus drives disease risk for wildlife, livestock, and humans. However, it remains challenging to quantify the availability of vertebrate hosts in field settings, particularly for medium-sized to large-bodied mammals. Here, we present a method that uses camera traps to quantify the availability of warm-bodied vertebrates to ticks. The approach is to deploy camera traps at questing height at a representative sample of random points across the study area, measure the average photographic capture rate for vertebrate species, and then correct these rates for the effective detection distance. The resulting "passage rate" is a standardized measure of the frequency at which vertebrates approach questing ticks, which we show is proportional to contact rate. A field test across twenty 1-ha forest plots in the Netherlands indicated that this method effectively captures differences in wildlife assemblage composition between sites. Also, the relative abundances of three life stages of the sheep tick Ixodes ricinus from drag sampling were correlated with passage rates of deer, which agrees with the known association with this group of host species, suggesting that passage rate effectively reflects the availability of medium- to large-sized hosts to ticks. This method will facilitate quantitative studies of the relationship between densities of questing ticks and the availability of different vertebrate species-wild as well as domesticated species-in natural and anthropogenic settings.

Cascading effects of predator activity on tick-borne disease risk.

Predators and competitors of vertebrates can in theory reduce the density of infected nymphs (DIN)-an often-used measure of tick-borne disease risk-by lowering the density of reservoir-competent hosts and/or the tick burden on reservoir-competent hosts. We investigated this possible indirect effect of predators by comparing data from 20 forest plots across the Netherlands that varied in predator abundance. In each plot, we measured the density of questing Ixodes ricinus nymphs (DON), DIN for three pathogens, rodent density, the tick burden on rodents and the activity of mammalian predators. We analysed whether rodent density and tick burden on rodents were correlated with predator activity, and how rodent density and tick burden predicted DON and DIN for the three pathogens. We found that larval burden on two rodent species decreased with activity of two predator species, while DON and DIN for all three pathogens increased with larval burden on rodents, as predicted. Path analyses supported an indirect negative correlation of activity of both predator species with DON and DIN. Our results suggest that predators can indeed lower the number of ticks feeding on reservoir-competent hosts, which implies that changes in predator abundance may have cascading effects on tick-borne disease risk.

New foci of Haemaphysalis punctata and Dermacentor reticulatus in the Netherlands.

In 2014 Haemaphysalis punctata was found in several locations on the mainland of the Netherlands for the first time since 1897. In the same areas Dermacentor reticulatus and Ixodes ricinus were found. Haemaphysalis punctata and D. reticulatus were tested for presence of Babesia spp. and Rickettsia spp. by PCR. Babesia spp. and spotted fever Rickettsiae were not detected in any of the collected H. punctata, while several D. reticulatus (6%) collected from the same areas were found to be positive for R. raoultii, a causative agent of tick-borne lymphadenopathy. We discuss the role of free-ranging domestic animals in maintaining H. punctata and D. reticulatus populations in dune areas in the Netherlands.