title: The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals.
abstract: Extant terrestrial biodiversity arguably is driven by the evolutionary success of angiosperm plants, but the evolutionary mechanisms and timescales of angiosperm-dependent radiations remain poorly understood. The Scarabaeoidea is a diverse lineage of predominantly plant- and dung-feeding beetles. Here, we present a phylogenetic analysis of Scarabaeoidea based on four DNA markers for a taxonomically comprehensive set of specimens and link it to recently described fossil evidence. The phylogeny strongly supports multiple origins of coprophagy, phytophagy and anthophagy. The ingroup-based fossil calibration of the tree widely confirmed a Jurassic origin of the Scarabaeoidea crown group. The crown groups of phytophagous lineages began to radiate first (Pleurostict scarabs: 108 Ma; Glaphyridae between 101 Ma), followed by the later diversification of coprophagous lineages (crown-group age Scarabaeinae: 76 Ma; Aphodiinae: 50 Ma). Pollen feeding arose even later, at maximally 62 Ma in the oldest anthophagous lineage. The clear time lag between the origins of herbivores and coprophages suggests an evolutionary path driven by the angiosperms that first favoured the herbivore fauna (mammals and insects) followed by the secondary radiation of the dung feeders. This finding makes it less likely that extant dung beetle lineages initially fed on dinosaur excrements, as often hypothesized.
authors.affiliation: Department of Life Sciences, Natural History Museum, London SW7 5BD, UK Zoologisches Forschungsmuseum Alexander Koenig Bonn, Adenauerallee 160, Bonn 53113, Germany ahrens.dirk_col@gmx.de.
authors.forename: Dirk
authors.initials: D
authors.lastname: Ahrens
authors.affiliation: Zoologisches Forschungsmuseum Alexander Koenig Bonn, Adenauerallee 160, Bonn 53113, Germany Division of Aquatic Ecology and Evolution, Institute of Ecology and Evolution, University of Bern, Bern, Switzerland EAWAG Centre of Ecology, Evolution and Biogeochemistry, Swiss Federal Institute for Aquatic Science and Technology, Kastanienbaum, Switzerland.
authors.forename: Julia
authors.initials: J
authors.lastname: Schwarzer
authors.affiliation: Department of Life Sciences, Natural History Museum, London SW7 5BD, UK Department of Life Sciences, Imperial College London, Silwood Park Campus, Ascot SL7 5PY, UK.
authors.forename: Alfried P
authors.initials: AP
authors.lastname: Vogler
mesh_terms: D000818:Animals; D044822:Biodiversity; D005075:Biological Evolution; D001517:Coleoptera / Q000235:genetics / Q000502:physiology*; D019143:Evolution, Molecular; D005247:Feeding Behavior; D005580:Fossils; D019476:Insect Proteins / Q000235:genetics; D019684:Magnoliopsida / Q000502:physiology; D008322:Mammals / Q000502:physiology; D010802:Phylogeny
chemical_list: D019476:Insect Proteins
keywords: Coleoptera; Scarabaeoidea; coprophagy; phytophagy; species diversity; time tree
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