Higher survival drives the success of nitrogen-fixing trees through succession in Costa Rican rainforests

Trees capable of symbiotic nitrogen (N) fixation ('N fixers') are abundant in many tropical forests. In temperate forests, it is well known that N fixers specialize in early-successional niches, but in tropical forests, successional trends of N-fixing species are poorly understood. We used a long-term census study (1997-2013) of regenerating lowland wet tropical forests in Costa Rica to document successional patterns of N fixers vs non-fixers, and used an individual-based model to determine the demographic drivers of these trends. N fixers increased in relative basal area during succession. In the youngest forests, N fixers grew 2.5 times faster, recruited at a similar rate and were 15 times less likely to die as non-fixers. As succession proceeded, the growth and survival disparities decreased, whereas N fixer recruitment decreased relative to non-fixers. According to our individual-based model, high survival was the dominant driver of the increase in basal area of N fixers. Our data suggest that N fixers are successful throughout secondary succession in tropical rainforests of north-east Costa Rica, and that attempts to understand this success should focus on tree survival.