Ant-induced evolutionary patterns in aphids

This review investigates ant-aphid mutualism (trophobiosis), in particular focusing on evolutionary processes in aphids resulting from this interaction. This broad literature review allows us to revise existing views on certain aspects of this mutualism and provide the first timeline of its possible development over a geological timescale. We propose a new classification of ant-aphid mutualism with respect to its ecological characteristics and present new explanations of the development of certain morphological structures of aphids as resulting from adaptation to a trophobiotic relationship with ants. In light of these findings, the presence of the so-called 'trophobiotic organ' in all myrmecophilous aphids is questioned. We review various communication modes between aphids and mutualistic ants and the possible influence of this symbiosis on modifications of aphid life cycles. Recent evidence on the suspected role of endosymbiotic bacteria in the development of mutualism is discussed. Finally, we propose a first timeline of ant-aphid interactions leading to the development of particular adaptations of aphids to mutualism, from general interactions between diverging Aphidomorpha and ants in the late Mesozoic to more specific coevolution between the aphid tribe Fordini and the ant genus Lasius.