BACKGROUND: The tribe Coccinellini is a group of relatively large ladybird beetles that exhibits remarkable morphological and biological diversity. Many species are aphidophagous, feeding as larvae and adults on aphids, but some species also feed on other hemipterous insects (i.e., heteropterans, psyllids, whiteflies), beetle and moth larvae, pollen, fungal spores, and even plant tissue. Several species are biological control agents or widespread invasive species (e.g., Harmonia axyridis (Pallas)). Despite the ecological importance of this tribe, relatively little is known about the phylogenetic relationships within it. The generic concepts within the tribe Coccinellini are unstable and do not reflect a natural classification, being largely based on regional revisions. This impedes the phylogenetic study of important traits of Coccinellidae at a global scale (e.g. the evolution of food preferences and biogeography). RESULTS: We present the most comprehensive phylogenetic analysis of Coccinellini to date, based on three nuclear and one mitochondrial gene sequences of 38 taxa, which represent all major Coccinellini lineages. The phylogenetic reconstruction supports the monophyly of Coccinellini and its sister group relationship to Chilocorini. Within Coccinellini, three major clades were recovered that do not correspond to any previously recognised divisions, questioning the traditional differentiation between Halyziini, Discotomini, Tytthaspidini, and Singhikaliini. Ancestral state reconstructions of food preferences and morphological characters support the idea of aphidophagy being the ancestral state in Coccinellini. This indicates a transition from putative obligate scale feeders, as seen in the closely related Chilocorini, to more agile general predators. CONCLUSIONS: Our results suggest that the classification of Coccinellini has been misled by convergence in morphological traits. The evolutionary history of Coccinellini has been very dynamic in respect to changes in host preferences, involving multiple independent host switches from different insect orders to fungal spores and plants tissues. General predation on ephemeral aphids might have created an opportunity to easily adapt to mixed or specialised diets (e.g. obligate mycophagy, herbivory, predation on various hemipteroids or larvae of leaf beetles (Chrysomelidae)). The generally long-lived adults of Coccinellini can consume pollen and floral nectars, thereby surviving periods of low prey frequency. This capacity might have played a central role in the diversification history of Coccinellini.

Australian National Insect Collection CSIRO GPO Box 1700 Canberra ACT 2601 Australia

Centre for Biodiversity Analysis Australian National University ACT Acton 2601 Australia

Centre for Molecular Biodiversity Research Museum Alexander Koenig Adenauerallee 53113 Bonn Germany

Centre for Molecular Biodiversity Research Museum Alexander Koenig Adenauerallee 53113 Bonn Germany Australian National Insect Collection CSIRO GPO Box 1700 Canberra ACT 2601 Australia

College of Environment and Plant Protection Hainan University No 58 Renmin Avenue Haikou 570228 China

Department of Evolutionary Biology and Ecology Institute of Biology 1 Albert Ludwig University of Freiburg Hauptstr 1 79104 Freiburg Germany

Institute of Entomology Biology Centre Branišovská 31 37005 České Budějovice CZ Czech Republic University of South Bohemia Branišovská 31 České Budějovice Czech Republic

Key Laboratory of Bio Pesticide Innovation and Application Guangdong Province Guangzhou China

Museum and Institute of Zoology Polish Academy of Sciences Wilcza 64 00 679 Warszawa Poland

State Key Laboratory of Biocontrol Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education Institute College of Ecology and Evolution Sun Yat Sen University Guangzhou 510275 China

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