title: 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.
abstract: 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.
authors.affiliation: Department of Biosciences, Centre for Ecological and Evolutionary Synthesis (CEES), University of Oslo, P.O. Box 1066 Blindern, NO-0316, Oslo, Norway. atle.mysterud@ibv.uio.no.
authors.forename: Atle
authors.initials: A
authors.lastname: Mysterud
authors.affiliation: Department of Biosciences, Centre for Ecological and Evolutionary Synthesis (CEES), University of Oslo, P.O. Box 1066 Blindern, NO-0316, Oslo, Norway. ragna.byrkjeland@gmail.com.
authors.forename: Ragna
authors.initials: R
authors.lastname: Byrkjeland
authors.affiliation: Department of Biosciences, Centre for Ecological and Evolutionary Synthesis (CEES), University of Oslo, P.O. Box 1066 Blindern, NO-0316, Oslo, Norway. lars.qviller@vetinst.no.|Norwegian Veterinary Institute, P.O. Box 750 Sentrum, NO-0106, Oslo, Norway. lars.qviller@vetinst.no.
authors.forename: Lars
authors.initials: L
authors.lastname: Qviller
authors.affiliation: Department of Biosciences, Centre for Ecological and Evolutionary Synthesis (CEES), University of Oslo, P.O. Box 1066 Blindern, NO-0316, Oslo, Norway. hildegunn.viljugrein@vetinst.no.|Norwegian Veterinary Institute, P.O. Box 750 Sentrum, NO-0106, Oslo, Norway. hildegunn.viljugrein@vetinst.no.
authors.forename: Hildegunn
authors.initials: H
authors.lastname: Viljugrein
mesh_terms: D000818:Animals; D049628:Body Size; D017753:Ecosystem; D065928:Forests; D018884:Ixodes / Q000254:growth & development*; D051379:Mice; D009664:Norway; D059208:Parasite Load; D012377:Rodentia / Q000469:parasitology*; D012621:Seasons; D012788:Shrews / Q000469:parasitology*; D013984:Tick Infestations / Q000453:epidemiology / Q000662:veterinary*