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Many aspects of the timeline and evolution of the dung beetles are still under debate. The age of origin of Scarabaeinae remains undetermined. Some phylogenetic analyses have proposed that the subfamily evolved in Africa during the Mesozoic (Ahrens et al. 2014; Tarasov & Génier 2015; Gunter et al. 2016) while others place its emergence in the early Cenozoic (Sole & Scholtz 2010; Mlambo et al. 2015).
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