Ticks prefer specific feeding sites on a host that are influenced by host–tick and tick–tick interactions. This study focused on the spatiotemporal distribution of ticks in Hokkaido sika deer, an important tick host in Hokkaido, Japan. Tick sampling was performed on the sika deer in the Shiretoko National Park between June and October 2022. Ticks were collected from 9 different body parts of the deer to compare their attachment site preferences. Interspecific and intraspecific relationships among ticks were examined using co-occurrence analysis. The collected ticks were nymphal and adult stages of 4 species: Ixodes ovatus, Ixodes persulcatus, Haemaphysalis japonica and Haemaphysalis megaspinosa. Seasonal variations in tick burden were observed, with I. persulcatus and I. ovatus peaking in June and declining towards October; H. japonica showing low numbers in July and August and increasing from September; and H. megaspinosa appearing from September onwards with little variation. Attachment site preferences varied among species, with a significant preference for the pinna in I. ovatus and I. persulcatus. Haemaphysalis japonica was mainly found on the body and legs between June and August, and shifted to the pinna from September. Haemaphysalis megaspinosa showed a general preference for areas other than the legs. Co-occurrence analysis revealed positive, negative and random co-occurrence patterns among the tick species. Ticks of the same genus and species exhibited positive co-occurrence patterns; I. ovatus showed negative co-occurrence patterns with Haemaphysalis spp. This study revealed the unique attachment site preferences and distinct seasonal distributions of tick species in the Hokkaido sika deer.

This review examines the phenomenon of co-feeding transmission in tick-borne pathogens. This mode of transmission is critical for the epidemiology of several tick-borne viruses but its importance for Borrelia burgdorferi sensu lato, the causative agents of Lyme borreliosis, is still controversial. The molecular mechanisms and ecological factors that facilitate co-feeding transmission are therefore examined with particular emphasis on Borrelia pathogens. Comparison of climate, tick ecology and experimental infection work suggests that co-feeding transmission is more important in European than North American systems of Lyme borreliosis, which potentially explains why this topic has gained more traction in the former continent than the latter. While new theory shows that co-feeding transmission makes a modest contribution to Borrelia fitness, recent experimental work has revealed new ecological contexts where natural selection might favour co-feeding transmission. In particular, co-feeding transmission might confer a fitness advantage in the Darwinian competition among strains in mixed infections. Future studies should investigate the ecological conditions that favour the evolution of this fascinating mode of transmission in tick-borne pathogens.

Experiments were conducted to investigate the potential for transmission of Orientia tsutsugamushi, the aetiological agent for scrub typhus, when naturally infected mite larvae were co-feeding with uninfected larvae. Larvae from colonies of Leptotrombidium deliense and L. imphalum infected with O. tsutsugamushi were used. Transmission of O. tsutsugamushi to previously uninfected L. deliense and Blankaartia acuscutellaris co-fed with infected L. deliense was shown to occur. The overall minimum rate of acquisition was 1·6% (4/258) for L. deliense and 2·5% (3/119) for B. acuscutellaris. When individual infected L. deliense were co-fed with B. acuscutellaris acquisition of O. tsutsugamushi was not detected. However, when 4 and 8 infected larvae were co-fed with B. acuscutellaris acquisition of O. tsutsugamushi was detected. Transmission of O. tsutsugamushi was not observed when uninfected L. deliense were co-fed with infected L. imphalum. This novel transmission route may explain the occurrence of rickettsiae in genera other than Leptotrombidium spp, which are considered to be the main vectors of O. tsutsugamushi.

We present observational and experimental evidence that cycles of the Lyme disease spirochaete, Borrelia burgdorferi s.l., can be maintained by sheep in the virtual absence of alternative hosts. A 2-year field study in upland moorland habitats of northwest UK established that sheep feed up to 80% of larval, >99% of nymphal and all of the adult female tick (Ixodes ricinus) population. Infection prevalence of B. burgdorferi in questing ticks reaches over 20%, but amplification of infection occurs principally as nymphs (20- to 30-fold), rather than larvae (4- to 7-fold), feed on sheep, and transmission from sheep to ticks occurred only during peak tick abundance in May and September. Experimental transmission studies confirmed that sheep, previously exposed to infected ticks on the moorland site, do not support systemic infections of B. burgdorferi, but they can transmit localized infections from infected to uninfected ticks co-feeding at the same site on the sheep's body.