5How symbionts are transmitted between hosts is key to determining whether sym-6 bioses evolve to be harmful or beneficial. Vertical transmission favors mutualistic sym-7 bionts, and horizontal transmission more virulent ones. Transmission mode evolution 8 itself depends on whether the host or symbiont can respond to selection on transmis-9 sion mode. When hosts control the transmission mode, vertical transmission should 10 evolve under more restrictive circumstances than when symbionts are in control. We 11 take a phylogenetic approach to determine whether the host, symbiont, or both control 12 transmission mode using the pooid grass-epichloid endophyte symbiosis as a model sys- 13 tem. This study is the first to investigate control of transmission mode evolution in a 14 phylogenetic context. We find a signal of host phylogeny but only in conjunction with 15 symbiont identity. This pattern suggests joint control of transmission mode by the host 16 and symbiont. It also suggests that non-genetic or non-conserved symbiont traits may 17 determine whether host traits lead to vertical or horizontal transmission. 18 1 19Symbiotic relationships are ubiquitous and can have large impacts on the fitness of the host, 20 symbiont, and organisms that interact with them [1,2]. Understanding the evolution of 21 symbiont virulence is therefore a matter of theoretical and practical interest. Transmission 22 mode is a key factor in virulence evolution. Vertical transmission favors mutualists, and 23 horizontal transmission parasites, assuming a positive relationship between virulence and 24 horizontal transmission [3,4] and in the absence of feedbacks selecting for mutualism [5, 6] 25 or parasitism [7]. Transmission mode evolution may itself depend on whether hosts or 26 symbionts can respond to selection on it [8]. We term this ability to respond to selection 27 "control," as selective pressures on the partner(s) in "control" determine the direction of 28 transmission mode evolution. For example, in the case of parasitism, symbiont control may 29 favor increased vertical transmission when host control does not. Despite its importance, 30 there has been little work exploring the patterns of transmission mode evolution over the 31 evolutionary history of extant symbioses.
32In this paper, we show that a phylogenetic perspective can provide valuable insight 33 into the control of transmission mode. In particular, we propose that if the variation in 34 transmission mode in a given symbiosis maps onto the phylogeny of one of the partners, we 35 can interpret it as that partner's traits determining the transmission mode, i.e., that partner 36 controls transmission mode. For example, if symbionts control transmission mode, related 37 symbiont species should be more likely to employ the same transmission mode than unrelated 38 species. If symbionts do not control transmission mode, then related symbionts should not 39 be more likely than unrelated symbionts to employ the same transmission mode, because 40 factors external to the sy...