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abdomen African species Afrotropical Afrotropical region arid associated Average length ball ball-rolling basal bidentate biological notes body shape Boheman Boucomont brood ovoids Cambefort Canthonini Cape carina Central Africa climatic clypeal clypeus coastal Colour convex cooler Coprini Copris Group d'Orbigny Diastellopalpus Dichotomiini distribution drier dry savanna dung beetles dung types East eastern Eastern Cape edge elephant dung elongate elytra endemic Epirinus Euoniticellus female finer-grained soils fore tibia forest genera genus genus comprises grassland habits Heliocopris herbivore highland istribution and biological Janssens Kheper kleptocoprid Klug large-bodied lineages lowland male margin Milichus moist savanna montane nesting behaviour occur omnivore dung Onitis Onthophagini Onthophagus Oriental region Pachysoma Palaearctic patterns Péringuey Phalops phylogeny Plate posterior primarily Proagoderus prothorax radiation rainfall region rainforest range recorded ridges Scarabaeini Scarabaeus Sceliages sexual dimorphism shrubland small-bodied South Africa southern Africa species groups species show sternites taxa tibiae tribe tropical tunnel Type-species upland West Africa whereas winter rainfall