On the difficult evolutionary transition from the free-living lifestyle to obligate symbiosis.

Obligate symbiosis evolved from free-living individuals most likely via the intermediate stage of facultative symbiosis. However, why should facultative symbionts, who can live independently but also benefit from their partners if these are available, give up this best of both worlds? Using the adaptive dynamics approach, we analyse a simple model, focusing on one partner of the symbiosis, to gain more insight into the selective forces that make individuals forgo the ability to reproduce in the free-living state. Our results suggest that, similar to the parasitism-mutualism continuum, the free-living way of life and obligate symbiosis are two extremes of a continuum of the ability to reproduce independently of a partner. More importantly, facultative symbiosis should be the rule as for many parameter combinations completely giving up independent reproduction or adopting a pure free-living strategy is not so easy. We also show that if host encounter comes at a cost, individuals that put more effort into increasing the chances to meet with their partners are more likely to give up the ability to reproduce independently. Finally, our model does not specify the ecological interactions between hosts and symbionts but we discuss briefly how the ecological nature of an interaction can influence the transition from facultative to obligate symbiosis.