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Article title
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Abstract
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Keywords
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Introduction
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Materials and methods
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Description of the morphological characters and their states
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Wing
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Epipharynx
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Labium
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Antennae
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Prothoracic apodemes
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Head
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Aedeagus and genital capsule
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Pygidium
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Abdominal ventrites
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Maxillae
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Pronota and prolegs
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Mandibles
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Elytra
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Cervical lateral sclerite
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Mesonotum
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Metanotum
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Meso- and metaventrite and meso- and metaleg
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Results
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Clades within the Oniticellini
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Clades within the Onthophagini
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Discussion
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The Oniticellini Clades
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Drepanocerina
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Helictopleurina
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Oniticellina
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The Tribe Onthophagini
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The Apical Clades of the Onthophagini
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The Genus Onthophagus
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Onthophagines
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Antennal Pits
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Nesting behavior in the dung feeders
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Inquiliny
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Necrophagy, mycetophagy, and frugivory
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Biogeography
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Conclusions
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Acknowledgments
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References
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