Background During the last decades a northward and upward range shift has been observed among many organisms across different taxa. In the northern hemisphere, ticks have been observed to have increased their latitudinal and altitudinal range limit. However, the elevational expansion at its northern distribution range remains largely unstudied. In this study we investigated the altitudinal distribution of the exophilic Ixodes ricinus and endophilic I. trianguliceps on two mountain slopes in Norway by assessing larval infestation rates on bank voles (Myodes glareolus). Methods During 2017 and 2018, 1325 bank voles were captured during the spring, summer and autumn at ten trapping stations ranging from 100 m to 1000 m.a.s.l. in two study areas in southern Norway. We used generalized logistic regression models to estimate the prevalence of infestation of both tick species along gradients of altitude, considering study area, collection year and season, temperature, humidity and altitude interactions as extrinsic variables, and host body mass and sex as intrinsic predictor variables. Results We found that both I. ricinus and I. trianguliceps infested bank voles at altitudes up to 1000 m.a.s.l., which is a substantial increase in altitude compared to previous findings for I. ricinus in this region. The infestation rates declined more rapidly with increasing altitude for I. ricinus compared to I. trianguliceps, indicating that the endophilic ecology of I. trianguliceps may provide shelter from limiting factors tied to altitude. Seasonal effects limited the occurrence of I. ricinus during autumn, but I. trianguliceps was found to infest rodents at all altitudes during all seasons of both years. Conclusions This study provides new insights into the altitudinal distribution of two tick species at their northern distribution range, one with the potential to transmit zoonotic pathogens to both humans and livestock. With warming temperatures predicted to increase, and especially so in the northern regions, the risk of tick-borne infections is likely to become a concern at increasingly higher altitudes in the future.

Rodents often act as important hosts for ticks and as pathogen reservoirs. At northern latitudes, rodents often undergo multi-annual population cycles, and the periodic absence of certain hosts may inhibit the survival and recruitment of ticks. We investigated the potential role of common shrews (Sorex araneus) to serve as a sup-How to cite this article: De Pelsmaeker, N., Korslund, L., & Steifetten, Ø. (2022). Host in reserve: The role of common shrews (Sorex araneus) as a supplementary source of tick hosts in small mammal communities influenced by rodent population cycles. Ecology and Evolution, 12, e8776.

In studies assessing tick abundance, the use of live traps to capture and euthanize rodent hosts is a commonly used method to determine their burden. However, captive animals can experience debilitating or fatal capture stress as a result prior to collection. An alternative method is the use of lethal traps, but this can potentially lead to tick drop-off between the time of capture and collection. In this study, in order to determine whether subjecting animals to capture stress is inevitable, we tested the difference in sheep tick (Ixodes ricinus) larval burdens between bank voles (Myodes glareolus) captured alive and euthanized, and lethally trapped bank voles. During 2017 and 2018, 1318 bank voles were captured using live (Ugglan Special no. 2) and lethal (Rapp2 Mousetrap) traps during two consecutive years over three seasons in two locations in Norway. Voles captured alive would remain captive until euthanized, while lethally trapped voles were killed instantly upon capture. Loglinear models, accounting for overdispersion, were used to determine whether trap type was influencing observed tick burden. Bank voles captured in lethal traps carried 5.7% more larvae compared to euthanized voles captured in live traps, but this difference was not significant (p = 0.420). Males were overall captured 2.7 times more frequently than females, and the sex ratio was equal in both trap types. This study shows that the use of lethal traps to determine tick burden of rodents is sufficiently reliable, without having to subject animals to potentially lethal stress, hereby reducing some ethical concerns of animal suffering and the results thereof, without compromising accuracy. Lethal trapping is also often more economical and practical, further favoring this collection method.

In recent decades, warming temperatures and changes in land use are supposed to have enabled several tick species to expand their distribution limit northwards. The progression of ticks to new areas may introduce new and emerging tick-borne pathogens as well as increase existing diseases. Aside from climatic conditions, ticks are dependent on hosts for survival, and rodents often act as important hosts for ticks and as pathogen reservoirs. At northern latitudes, rodents often undergo multi-annual population cycles, and the periodic absence of hosts may inhibit the further progression of ticks. We investigated the potential role of common shrews (Sorex araneus) to serve as a stable host source to immature life stages of a generalist tick Ixodes ricinus and a specialist tick I. trianguliceps, during decreasing abundances of bank voles (Myodes glareolus). We tested whether ticks would have a propensity to parasitize a certain host type dependent on host population size and composition in two high latitude locations in southern Norway, by comparing tick burdens on trapped animals. We found that I. ricinus larvae showed an equal propensity to parasitize both host types as the host population composition changed, but voles had a higher level of parasitism by nymphs (p< 0.004). An overall larger host population size favored the parasitism of voles by larvae (p= 0.027), but not by nymphs (p= 0.074). I. trianguliceps larvae showed a higher propensity to parasitize shrews, regardless of host population size or composition (p= 0.004), while nymphs parasitized shrews more as vole abundance increased (p= 0.002). The results indicate that common shrews may have the potential to act as a replacement host during periods of low rodent availability, but long-term observations encompassing complete rodent cycles may determine whether shrews are able to maintain tick range expansion despite low rodent availability.

Background: During the last decades a northward and upward range shift has been observed among many organisms across different taxa. In the northern hemisphere, ticks have been observed to have increased their latitudinal and altitudinal range limit. However, the elevational expansion at its northern distribution limit remains largely unstudied. In this study we investigated the altitudinal distribution of the exophilic Ixodes ricinus and endophilic I. trianguliceps on two mountain slopes in Norway by assessing larval infestation rates on bank voles (Myodes glareolus).Methods: During 2017 and 2018, 1325 bank voles were captured during spring, summer and autumn at 10 trapping stations ranging from 100 m to 1000 m.a.s.l in two study areas in southern Norway. We used generalized logistic regression models to estimate the prevalence of infestation of both tick species along altitude, considering study area, collection year and season, temperature, humidity and altitude interactions as extrinsic variables; and host body mass and sex as intrinsic predictor variables.Results: We found that both I. ricinus and I. trianguliceps infested bank voles at altitudes up to 1000 m.a.s.l., which is a substantial increase in altitude compared to previous findings for I. ricinus in this region. The infestation rates declined more rapidly for I. ricinus compared to I. trianguliceps, indicating that the endophilic ecology of I. trianguliceps may provide shelter from limiting factors tied to altitude. Seasonal effects limited the occurrence of I. ricinus during autumn, but I. trianguliceps was found to infest rodents at all altitudes during all seasons of both years.Conclusions: This study provides new insights into the altitudinal distribution of two tick species at their northern distributional limit, one with the potential to transmit zoonotic pathogens to both humans and livestock. With warming temperatures predicted to increase, and especially so in the northern regions, the risk of tick-borne infections is likely to become a concern at increasingly higher altitudes in the future.