An updated meta-analysis of the distribution and prevalence of Borrelia burgdorferi s.l. in ticks in Europe

Background: The bacteria of the group Borrelia burgdorferi s.l. are the etiological agents of Lyme borreliosis in humans, transmitted by bites of ticks. Improvement of control measures requires a solid framework of the environmental traits driving its prevalence in ticks. Methods: We updated a previous meta-analysis of the reported prevalence of Borrelia burgdorferi s.l. in questing nymphs of Ixodes ricinus with a literature search from January 2010-June 2017. This resulted in 195 new papers providing the prevalence of Bb for 926 geo-referenced records. Previously obtained data (878 records, years 2000-2010) were appended for modelling. The complete dataset contains data from 82,004 questing nymphs, resulting in 558 records of B. afzelii, 404 of B. burgdorferi s.s. (only 80 after the year 2010), 552 of B. garinii, 78 of B. lusitaniae, 61 of B. spielmanii, and 373 of B. valaisiana. We associated the records with explicit coordinates to environmental conditions and to a categorical definition of European landscapes (LANMAP2) looking for a precise definition of the environmental niche of the most reported species of the pathogen, using models based on different classification methods. Results: The most commonly reported species are B. afzelii, B. garinii and B. valaisiana largely overlapping across Europe. Prevalence in ticks is associated with portions of the environmental niche. Highest prevalence occurs in areas of 280 degrees-290 degrees (Kelvin) of mean annual temperature experiencing a small amplitude, steady spring slope, together with high mean values and a moderate spring rise of vegetation vigor. Low prevalence occurs in sites with low and a noteworthy annual amplitude of temperature and the Normalized Difference Vegetation Index (colder areas with abrupt annual changes of vegetation). Models based on support vector machines provided a correct classification rate of the habitat and prevalence of 89.5%. These results confirm the association of prevalence of the three most commonly reported species of B. burgdorferi s.l. in Europe to parts of the environmental niche and provide a statistically tractable framework for analyzing trends under scenarios of climate change.