We used entire mitochondrial (mt) genome sequences (14.5–15 kbp) to resolve the phylogeny of the four main lineages of the Haematobothrion ticks: Alloceraea, Archaeocroton, Bothriocroton and Haemaphysalis . In our phylogenetic trees, Alloceraea was the sister to Archaeocroton sphenodonti , a tick of an archetypal reptile, the tuatara, from New Zealand, to the exclusion of the rest of the species of Haemaphysalis . The mt genomes of all four of the Alloceraea species that have been sequenced so far had a substantial insert, 132–312 bp, between the tRNA‐Glu (E) gene and the nad1 gene in their mt genomes. This insert was not found in any of the other eight subgenera of Haemaphysalis . The mt genomes of 13 species of Haemaphysalis from NCBI GenBank were added to the most recent data set on Haemaphysalis and its close relatives to help resolve the phylogeny of Haemaphysalis , including five new subgenera of Haemaphysalis not previously considered by other authors: Allophysalis (structurally primitive), Aboimisalis (structurally primitive), Herpetobia (structurally intermediate), Ornithophysalis (structurally advanced) and Segalia (structurally advanced). We elevated Alloceraea Schulze, 1919 to the status of genus because Alloceraea Schulze, 1919 is phylogenetically distinct from the other subgenera of Haemaphysalis . Moreover, we propose that the subgenus Allophysalis is the sister to the rest of the Haemaphysalis (14 subgenera) and that the ‘structurally primitive’ subgenera Hoogstraal and Kim comprise early diverging lineages. Our matrices of the pairwise genetic difference (percent) of mt genomes and partial 16S rRNA sequences indicated that the mt genome sequence of Al . kitaokai (gb# OM368280) may not be Al . kitaokai Hoogstraal, 1969 but rather another species of Alloceraea . In a similar way, the mt genome sequence of H . (Herpetobia ) nepalensis Hoogstraal, 1962 (gb# NC_064124) was only 2% genetically different to that of H . ( Allophysalis ) tibetensis Hoogstraal, 1965 (gb# OM368293): this indicates to us that they are the same species. Alloceraea cretacea may be better placed in a genus other than Alloceraea Schulze, 1919. Reptiles may have been the host to the most recent common ancestor of Archaeocroton and Alloceraea .

Mitochondrial (mt) genomes have played a major role in elucidating evolutionary relationships in ticks (Ixodida), especially in soft ticks (Argasidae). Mitochondrial genomes have been sequenced for representatives of most of the genera and other major lineages. This includes members of the Afrotropical, Nearctic, Neotropical and Palearctic Pavlovskyella Pospelova-Shtrom, 1950, which is a subgenus of Ornithodoros Koch, 1844. These continent-associated lineages do not form a monophyletic group: rather, the subgenus Pavlovskyella is paraphyletic. The only zoogeographic region for which mt genomes are not available is the Australasian region. Here we report the first mt genome of an Australasian Pavlovskyella: Ornithodoros (Pavlovskyella) gurneyi Warburton, 1926, the kangaroo soft tick. [The companion paper to the present work, Barker et al. (2025), presents the mt genome, 18S and 28S rRNA of the only other known Australasian Pavlovskyella, McMillan’s Australian tree-hollow argasid, Ornithodoros (Pavlovskyella) macmillani Hoogstraal & Kohls, 1966.] Our phylogenetic trees reveal that the two Australasian Pavlovskyella have their own clade (i.e. are sister-taxa) which does not have a sister-group relationship with species of Pavlovskyella from any of the four other zoogeographic regions. We propose that the Pavlovskyella from the Afrotropical, Australasian, Nearctic, Neotropical and Palearctic zoogeographic regions have independent evolutionary histories and thus are best considered different genera. Molecular dating leads us to propose that continental drift may have caused the evolution of five different lineages of Pavlovskyella, each in a different zoogeographic region of the world.

Revealing interactions between ticks and wild animals is vital for gaining insights into the dynamics of tick-borne pathogens in the natural environment. We aimed to elucidate the factors that determine tick infestation in wild animals by investigating ticks on invasive raccoons (Procyon lotor) in Hokkaido, Japan. We first examined the composition, intensity, and seasonal variation of ticks infesting raccoons in six study areas in Hokkaido from March 2022 to August 2023. In one study area, ticks infesting tanukis (raccoon dog, Nyctereutes procyonoides albus) were collected in May to July in both 2022 and 2023, and questing ticks were collected from the vegetation by flagging every other week in the same period. Next, we screened 17 environmental and host variables to determine factors that affect the number of ticks infesting raccoons using generalized linear (mixed) models. From 245 raccoons, we identified a total of 3,917 ticks belonging to eight species of two genera: the most prominent species were Ixodes ovatus (52.9 %), followed by Haemaphysalis megaspinosa (14.4 %), Ixodes tanuki (10.6 %), and Ixodes persulcatus (9.5 %). Ixodes ovatus was also predominant among questing ticks and ticks infesting tanukis. Although I. tanuki was frequently collected from raccoons and tanukis, it was rarely collected in the field. The variables that significantly affected the infestation on raccoons differed by genus, species and developmental stage of the tick. For instance, the infestation of adult I. ovatus was significantly affected by four variables: night-time temperature during nine days before capturing the raccoon, the size of forest area around the capture site, sex of the raccoon, and sampling season. The first two variables were also responsible for the infestation on raccoons of almost all species and stages of ticks. Our study revealed that the number and composition of ticks infesting raccoons can be affected not only by landscape of their habitats but also by weather conditions in several days before capturing.