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Background  In Europe, hard ticks of the subgenus  Pholeoixodes  (Ixodidae:  Ixodes ) are usually associated with burrow-dwelling mammals and terrestrial birds. Reports of  Pholeoixodes  spp. from carnivores are frequently contradictory, and their identification is not based on key diagnostic characters. Therefore, the aims of the present study were to identify ticks collected from dogs, foxes and badgers in several European countries, and to reassess their systematic status with molecular analyses using two mitochondrial markers.    Results  Between 2003 and 2017, 144  Pholeoixodes  spp. ticks were collected in nine European countries. From accurate descriptions and comparison with type-materials, a simple illustrated identification key was compiled for adult females, by focusing on the shape of the anterior surface of basis capituli. Based on this key, 71 female ticks were identified as  I. canisuga , 21 as  I. kaiseri  and 21 as  I. hexagonus . DNA was extracted from these 113 female ticks, and from further 31 specimens. Fragments of two mitochondrial genes,  cox 1 (cytochrome  c  oxidase subunit 1) and 16S rRNA, were amplified and sequenced.  Ixodes kaiseri  had nine unique  cox 1 haplotypes, which showed 99.2–100% sequence identity, whereas  I. canisuga  and  I. hexagonus  had eleven and five  cox 1 haplotypes, respectively, with 99.5–100% sequence identity. The distribution of  cox 1 haplotypes reflected a geographical pattern.  Pholeoixodes  spp. ticks had fewer 16S rRNA haplotypes, with a lower degree of intraspecific divergence (99.5–100% sequence identity) and no geographical clustering. Phylogenetic analyses were in agreement with morphology:  I. kaiseri  and  I. hexagonus  (with the similar shape of the anterior surface of basis capituli) were genetically more closely related to each other than to  I. canisuga . Phylogenetic analyses also showed that the subgenus  Eschatocephalus  (bat ticks) clustered within the subgenus  Pholeoixodes .    Conclusions  A simple, illustrated identification key is provided for female  Pholeoixodes  ticks of carnivores (including  I. hexagonus  and  I. rugicollis ) to prevent future misidentification of these species. It is also shown that  I. kaiseri  is more widespread in Europe than previously thought. Phylogenetic analyses suggest that the subgenus  Pholeoixodes  is not monophyletic: either the subgenus  Eschatocephalus  should be included in  Pholeoixodes , or the latter subgenus should be divided, which is a task for future studies.

parasites and vectors

Contributions to the phylogeny of Ixodes (Pholeoixodes) canisuga, I. (Ph.) kaiseri, I. (Ph.) hexagonus and a simple pictorial key for the identification of their females