Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae) Génier, François Moretto, Philippe Zootaxa 2017 4248 1 1 110 363WG [151,475,1519,1545] Insecta Scarabaeidae Digitonthophagus Animalia Coleoptera 14 15 Arthropoda species bonasus   Diagnosis. Male cephalic horns long, obliquely divergent in frontal view ( Figs. 24, 26, 28); females with horns on vertex ( Figs. 71–73).