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abdomen African species Afrotropical Afrotropical region annual rainfall arid associated Average length ball ball-rolling basal bidentate biological notes body shape Boheman Boucomont brood ovoids Cambefort Canthonini Cape carina Central Africa climate clypeal clypeus coastal Colour cooler Coprini Copris Group d'Orbigny Diastellopalpus Dichotomiini Distribution and biological drier dry savanna dung beetles dung types East eastern Eastern Cape edge elephant dung elongate elytra endemic eneric characters Epirinus Euoniticellus female finer-grained soils fore tibia forest genera genus genus comprises grassland habits Heliocopris herbivore highland Janssens Kheper kleptocoprid large-bodied lineages male margin Milichus moist savanna montane nesting behaviour occur omnivore dung Onitis Onthophagus Oriental region Pachysoma Palaearctic patterns Peringuey Phalops phylogeny Plate posterior primarily Proagoderus prothorax radiation rainfall region rainforest range recorded ridges Scarabaeini Scarabaeus Sceliages Scholtz and Howden 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