title: Mandible evolution in the Scarabaeinae (Coleoptera: Scarabaeidae) and adaptations to coprophagous habits.
abstract: BACKGROUND
  The astonishing spectrum of scarabaeine lifestyles makes them an attractive group for studies in entomology and evolutionary biology. As a result of adaptions to specific food substrates and textures, the mouthparts of dung beetles, particularly the mandible, have undergone considerable evolutionary changes and differ distinctly from the presumptive ancestral conditions of the Coleoptera and Polyphaga. The possible functions of dung beetle mouthparts and the evolution of dung feeding have been controversial for decades.
  
  
  RESULTS
  In this study, 187 scarabs representing all tribes of the Scarabaeinae and the major lineages within the Scarabaeoidea, along with three major feeding types within the Scarabaeoidea (omnivory, phytophagy and coprophagy), were studied. Based on geometric morphometric and three-dimensional (3D) reconstruction approaches, morphological differences in mandibles among the three feeding types were identified. The ancestral forms of the mandible within the Scarabaeinae were reconstructed and compared with those of modern species. The most recent common ancestor of the Scarabaeinae fed on soft materials, and the ancestor of the Scarabaeinae and the Aphodiinae was in an evolutionary transition between processing more solid and softer substrates.
  
  
  CONCLUSIONS
  Coprophagy originated from omnivorous ancestors that were very likely saprophagous. Furthermore, phytophagy may also have originated from omnivory. In addition, our study addresses the integration and modularity of geometric morphometric data in a phylogenetic context.
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.
authors.forename: Ming
authors.initials: M
authors.lastname: Bai
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.|University of Chinese Academy of Sciences, Yuquan Road, Shijingshan Beijing, 100039 P. R. China.
authors.forename: Sha
authors.initials: S
authors.lastname: Li
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.|University of Chinese Academy of Sciences, Yuquan Road, Shijingshan Beijing, 100039 P. R. China.
authors.forename: Yuanyuan
authors.initials: Y
authors.lastname: Lu
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.
authors.forename: Haidong
authors.initials: H
authors.lastname: Yang
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.
authors.forename: Yijie
authors.initials: Y
authors.lastname: Tong
authors.affiliation: Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District Beijing, 100101 People's Republic Of China.
authors.forename: Xingke
authors.initials: X
authors.lastname: Yang
keywords: 3D; Coprophagy; Dung beetle; Geometric morphometric; Mouthparts