Redescription of Alloceraea cretacea (Acari: Ixodida) with an additional nymphal fossil added to this species: novel insights into the evolution of the Haematobothrion

Abstract A fossil assigned to the extant ixodid genus Haemaphysalis was previously described from the Late Cretaceous (ca. 99 Ma) Burmese amber. Haemaphysalis ( Alloceraea ) cretacea was considered the oldest, and only, fossil representative of this genus. Significant criticism was raised regarding possible misidentification of this amber fossil. Microtomography of the original holotype and new material allows new perspectives on this controversy. A new fossil nymph is described and both fossils are considered Alloceraea cretacea comb. nov. nymphs based on a series of morphological characters: no genital aperture, eyeless, 11 festoons, coxa I simple with a short, wide triangular spur. Specific morphological features for the ‘structurally primitive’ Alloceraea are discussed and include palpi elongate, with long setae on palps, the hypostome dorsally longer than the chelicera and the corona visible distally and with a specific distribution of the denticles. Alloceraea and its sister genus Archaeocroton (that includes amber fossil taxa), share a common ancestor with Haemaphysalis sensu stricto, indicating a minimum divergence time of at least 100 MYA for these lineages. In addition, a fossil of Bothriocroton has also been described from Burmese amber. This genus with Cryptocroton groups basal to the Alloceraea / Archaecroton-Haemaphysalis assemblage forming a monophyletic clade named Haematobothrion. A synthesis of the fossil information and the current systematic understanding of the Haematobothrion allows new hypotheses about the origins of this group and the various lineages it comprises. Notably that the major Haematobothrion lineages originated and diverged during the time period when the Burma terrane was migrating from Australia to Asia.