“…Previous studies on microbes within historical bone tissue mainly focussed on the remnants of microbial activity [32,36], the pathologic lesions caused by microbial infection [68,83], or the identification of ancient DNA (aDNA) from pathogens [30,33,58]. However, recent studies on histopathology of historical dry bone provide organic, more 1 3 precisely cellular, bacterial and fungal evidence [16,17,64,71]. With regard to the treponematoses, molecular and microscopic diagnosis in human skeletal remains continues to be challenging [2,86].…”