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Flea beetles (Alticini) are a diverse lineage of leaf beetles (Coleoptera: Chrysomelidae) with the majority of genera living on vascular plants. However, several genera are associated with mosses or leaf litter, as an alternative strategy, often accompanied by changes in morphology and biology. Here, we present the first comprehensive DNA-based phylogenetic analysis of Alticini including 13 moss-inhabiting genera. We show that moss-association is a common phenomenon in Alticini; based on our ancestral-state mapping, we reveal at least nine independent origins of moss-inhabiting, and four independent origins of leaf-litter-inhabiting life-strategy. Our findings illustrate that both cryptic strategies induce parallel evolution of similar ecomorphological adaptations, such as flight loss and a convex body shape. An overview of known moss-inhabiting flea beetle genera is also provided. We describe Benedictoides munclingeri gen. et sp. nov. from the cloud forests of Mount Cameroon, the first flea beetle genus clearly assigned to the moss-inhabiting strategy from tropical Africa. Additionally, our phylogeny provides new insights into the relationships among various flea beetle genera. We estimate six main clades grouping the majority of studied genera, and propose several new generic groups. Along with our phylogenetic results, we discuss moss-inhabiting flea beetle evolution and perspectives of their systematics. [ABSTRACT FROM AUTHOR]