What do we still need to know about Ixodes ricinus?

In spite of many decades of intensive research on Ixodes ricinus, the castor bean tick of Europe, several important aspects of its basic biology remain elusive, such as the factors determining seasonal development, tick abundance and host specificity, and the importance of water management. Additionally, there are more recent questions about the geographical diversity of tick genotypes and phenotypes, the role of migratory birds in the ecoepidemiology of I. ricinus, the importance of protective immune responses against I. ricinus, particularly in the context of vaccination, and the role of the microbiome in pathogen transmission. Without more detailed knowledge of these issues, it is difficult to assess the likely effects of changes in climate and biodiversity on tick distribution and activity, to predict potential risks arising from new and established tick populations and I. ricinus-borne pathogens, and to improve prevention and control measures. This review aims to discuss the most important outstanding questions against the backdrop of the current state of knowledge of this important tick species.