Lachnine aphids are unusual among phytophagous insects because they feed on both leafy and woody parts of both angiosperm and conifer hosts. Despite being piercing-sucking phloem-feeders, these aphids are most speciose on woody parts of coniferous hosts. To evaluate the significance of this unusual biology on their evolution, we reconstructed the ancestral host and feeding site of the lachnine aphids and estimated important host shifts during their evolution. We sampled 78 species representing 14 of the 18 genera of Lachninae from Asia and North America. We performed parsimony, Bayesian and likelihood phylogenetic analyses of combined mitochondrial Cox1, Cox2, CytB and nuclear EF1a1 DNA sequences. We dated the resulting phylogram's important nodes using Bayesian methods and multiple fossil and secondary calibrations. Finally, we used parsimony and Bayesian ancestral state reconstruction to evaluate ancestral feeding ecology. Our results suggest the lachnine common ancestor fed on a woody part of an angiosperm host in the mid-Cretaceous. A shift to conifer hosts in the Late Cretaceous is correlated with a subsequent increased diversification in the Palaeogene, but a switch to leafy host tissues did not engender a similar burst of diversification. Extant lachnine lineages exhibit the full range of historical association with their hosts: some appeared before, some concomitant with and some after the appearance of their hosts. We conclude our study by placing all the lachnine genera in five tribes.

• 31430078/National Natural Sciences Foundation of China
• 31372237/National Natural Sciences Foundation of China
• 2011FY120200/Ministry of Science and Technology of the People's Republic of China
• 2014FY210200/Ministry of Science and Technology of the People's Republic of China
• 2013FY111200/Ministry of Science and Technology of the People's Republic of China

• XDA05080700/"Strategic Priority Research Program" of the Chinese Academy of Sciences
• J1210002/National Science Fund for Fostering Talents in Basic Research