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RESEARCH LETTER a N122H Danaus plexippus A119 Danaus erippus Danaus chrysippus L111V Danaus gilippus N122 Feeding Danaus genutia Sequestration Tirumala septentrionis Tirumala limniace Tirumala petiveran Parantica aglea leopsis juventa Amauris tartarea Euploea phaenareta A119S Euploea mulciber Q111L Euploea core godartii ie Idea leuconoe Lycorea halia Mechanitis polymnia 12 13 14 15 Heliconius erato 19 Drosophila melanogaster cliviconlineata versicolora 10 100 7 hadnus latus Camponotus floridanus Apis mellifera esis rapedata 60 Pediculusingtu's m U Q W W D Ou trhocoris apt Larval to adult survival (%) No change Lygaous kalmil B 40 - vomorpha halys Feeding 111 or 119 ASR ASP lectularius Sequestration 111 + 119 or 119 + 122 20 - siphon pisur 111 + 119 + 122 on O 7.5- Independent origins of specialization 9.0 8 10.5- 12.0 xxx 1 2 3 4 5 6 1 8 9 10 1 12 13 14 15 16 17 18 19 20 21 d Quabain (mM) Fig. 2 | Drosophila flies with edited genomes path of ATPo in the monarch butterfly lineag survival when fed cardiac glycosides. a, The m Frequency Time ENY QSN QNN QSN LSN QAN UNN LSN QSN SN QSN LAN with the substitutions observed in the HI-H2 1 2 20- Petschenka et al.) . b, Non-synonymous point TSH ESN ONN TNH QSY UNH USH LSN QSN LSN QSN LSN ASH QAN LSN LAN LAN LSN DNA sequence of the native Atpa in Drosophila the substitutions at sites 111, 119 and 122. Codo 0.2 ISN LSN QSH LSN VSN c, d, Larval-adult survival (c) and adult survival P value diets with ouabain were different between mon aSH USH and control lines (QAN = engineered control; Symbols represent the mean I s.e.m. of 3-6 bio Fig. 1 | Mutational paths in ATPo associated with insect specialization sequestering), and the number of substituted sites at positions 111, 119 (50 larvae and 10 females per replicate in c and on cardiac glycoside-producing plants are constrained. a, Protein and 122 along branches in grey-scale (light grey 0, medium grey 1, dark were fit using a logistic regression model for ead homology model of Drosophila melanogaster ATPa (navy) superimposed grey 2, black 3), based on maximum likelihood ASR of H1-H2 loo in survivorship trajectories between lines were on a Sus scrofa ATPa crystal structure (light grey) with ouabain (yellow) aminoa sequences. Black asterisks indi X the Atpa copy number for in the binding pocket. Residues 111, 119 and 122 (sticks) within the species with multiple paralogues. c, ATPo substitutions inferred from ASR HI-H2 extracellular loop are associated with feeding on cardiac at p ed) in 21 lineages where butterfly, suggesting that TSI is the predomin glycoside-producing plants and toxin sequestration. b, Maximum specialization occurred independently. d, P value distribution from a for the in vivo toxin resistance observed abo