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prokary-oticcellsand10–100timesmoreviruses!),and genetically extremely diversied. Sucha large assemblage cannot be ignoredin attempts to understand life’s historyon Earth, however, how can biologistsaccount for its evolution?For long, the notion of descent withmodication, describing a process ofvertical inheritance, dening a tree-likegenealogical pattern, when the geneticmaterial, modied by some mutations, istransferred from the genome of a lastcommon ancestor to its direct progenyhas offered a promising way to classifyorganisms and species. Thus, the origin ofmicrobial adaptations can be searched forwithin lineages, in the changes of geneticmaterial inherited from one ancestor. Yet,such studies are strongly constrained.Non-treelikeevolution,generatingaretic-ulate evolutionary pattern, cannot be ana-lyzed with a genealogical tree. Moreover,viruses do not all originate from a singlelast ancestor (Lima-Mendez et al., 2008),nor do they all display obvious genealog-ical relationships with cellular organisms,hindering the collective study of mobilegenetic elements and cells with a singletree.Genealogy also plays a central rolefor explaining the main types of behav-iors described in the biological world:selshness, mutualism, altruism, and spite(West et al., 2006). The evolution ofthese interactions can be understood byaccounting for kinship between protag-onists, under the standard assumptionthat genealogical proximity between indi-viduals entails their genetic proximity(Huneman, 2013). Thus, knowing the rel-ative kinship, the benet for the recipientof an interaction and the cost for its actorallows determining when an individualcooperating with a kin, in ways enhanc-ing its reproduction, or preventing dis-tantly related members of a population toreproduce, actually maximizes the repro-ductive success and the survival of its owngenes (van Baalen, 2013). Microbiologistsarethusinclinedtoembracetheconcep-tual framework of kin selection to analyzemany cooperations (Diggle et al., 2007).However, it is probably not enoughto take genealogical relationships intoaccount to explain the diversity, evolution,andinteractionsinthemicrobialworld.Toosystematicafocusongenealogymayevenintroducesomebiasesintheexplana-tions of microbial diversity, evolution andinteractions, because many crucial bio-logical phenomena result from processesorthogonal to vertical descent.

frontiers in microbiology

The origins of microbial adaptations: how introgressive descent, egalitarian evolutionary transitions and expanded kin selection shape the network of life