Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants.

African Butterfly Research Institute Nairobi Kenya

Animal Biology Division Institute of Biological Sciences University of the Philippines Los Baños Laguna Philippines

Aquapro Villavicencio Colombia

Australian Museum Sydney New South Wales Australia

Australian National Insect Collection Canberra Australian Capital Territory Australia

Biology Centre CAS České Budějovice Czech Republic

Biology Department City College of New York City University of New York New York NY USA

Botany and National Herbarium Division National Museum of the Philippines Manila Philippines

Center for Biodiversity and Global Change Yale University New Haven CT USA

Center for Biodiversity Research Department of Biological Sciences University of Memphis Memphis TN USA

Center for Global Mountain Biodiversity Globe Institute University of Copenhagen Copenhagen Denmark

Center for Integrative Conservation Xishuangbanna Tropical Botanical Garden Chinese Academy of Sciences Mengla China

Center for Macroecology Evolution and Climate GLOBE Institute University of Copenhagen Copenhagen Denmark

Centro de Ciências Naturais e Exatas Pós Graduação em Biodiversidade Animal Universidade Federal de Santa Maria Santa Maria Brazil

Chiang Mai Thailand

CNRS Institut des Sciences de l'Evolution de Montpellier Montpellier France

College of Arts and Sciences Jose Rizal Memorial State University Tampilisan Philippines

College of Life Science National Taiwan Normal University Taipei Taiwan

Departamento de Zoologia Universidade Federal do Rio Grande do Sul Porto Alegre Brazil

Department of Biology Georgetown University Washington DC USA

Department of Biology University of Florida Gainesville FL USA

Department of Biology University of Pennsylvania Philadelphia PA USA

Department of Biology Washington University in St Louis St Louis MO USA

Department of Botany and Plant Sciences University of California Riverside Riverside CA USA

Department of Ecology and Evolutionary Biology Yale University New Haven CT USA

Department of Ecology Swedish University of Agricultural Sciences Uppsala Sweden

Department of Entomology and Plant Pathology Auburn University Auburn AL USA

Department of Entomology College of Plant Protection South China Agricultural University Guangzhou China

Department of Entomology National Museum of Natural History Smithsonian Institution Washington DC USA

Department of Entomology Natural History Museum of Geneva Geneva Switzerland

Department of Evolution Ecology and Organismal Biology The Ohio State University Columbus OH USA

Department of Karyosystematics Zoological Institute of Russian Academy of Sciences St Petersburg Russia

Department of Life and Earth Sciences Perimeter College Georgia State University Decatur GA USA

Department of Organismic and Evolutionary Biology and Museum of Comparative Zoology Harvard University Cambridge MA USA

Department of Plant and Wildlife Sciences Brigham Young University Provo UT USA

Division of Ecology and Evolution Research School of Biology The Australian National University Acton Canberra Australian Capital Territory Australia

Entomology and Nematology Department University of Florida Gainesville FL USA

Entomology Laboratory Museo de Zoología Universidad del Azuay Cuenca Ecuador

Entomology Section National Museum of Natural History Manila Philippines

Faculty of Science Department of Zoology University of South Bohemia České Budějovice Czech Republic

Florida Museum of Natural History University of Florida Gainesville FL USA

Hachiôji Japan

Institut Botànic de Barcelona Barcelona Spain

Institut de Biologia Evolutiva Barcelona Spain

Institute of Silviculture and Forest Protection University of West Hungary Sopron Hungary

Iziko South African Museum Cape Town South Africa

Köln Germany

Laboratorio de Ecología y Morfometría Evolutiva Centro de Investigación de Estudios Avanzados del Maule Universidad Católica del Maule Talca Chile

Leibniz Institute for the Analysis of Biodiversity Change Zoological Research Museum Alexander Koenig Bonn Germany

McGuire Center for Lepidoptera and Biodiversity Florida Museum of Natural History University of Florida Gainesville FL USA

Millennium Institute Biodiversity of Antarctic and Subantarctic Ecosystems Santiago Chile

Museo de Historia Natural Universidad Nacional Mayor de San Marcos Lima Peru

National Centre for Biological Sciences Tata Institute of Fundamental Research Bengaluru India

Natural History Museum London UK

PhD Program in Biology Graduate Center City University of New York New York NY USA

RAPiD Genomics Gainesville FL USA

Research Center for Biosystematics and Evolution National Research and Innovation Agency Cibinong Bogor Indonesia

T G Masaryk Water Research Institute Prague Czech Republic

The Research Institute of Evolutionary Biology Setagaya Japan

The University Museum The University of Tokyo Tokyo Japan

Translational Data Analytics Institute The Ohio State University Columbus OH USA

Turkana Basin Institute Stony Brook University Stony Brook NY USA

University of Chinese Academy of Sciences Beijing China

Vietnam National Museum of Nature Vietnam Academy of Science and Technology Hanoi Vietnam

Vietnam Programme Fauna and Flora International Hanoi Vietnam

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