Contrasting effects of nutrients and consumers on tree colonization and growth during secondary succession

For succession to proceed from herbaceous to woody dominance, trees must colonize herbaceous communities and grow. Success across these two phases of succession might result from different interactions with the herbaceous community. First, colonizing trees must compete against larger, established herbs, while subsequent growth occurs among similarly sized or smaller herbs. This shift from colonization to growth may cause three drivers of secondary succession-- nutrients, consumers, and herbaceous diversity--to differentially affect tree colonization and growth. Initially, these drivers should favor larger, established herbs, reducing colonization. Later, when established trees can better compete with herbs, these drivers should benefit trees and increase their growth. In a four-year study, we added nutrients to, excluded aboveground consumers from, and manipulated initial richness of, the herbaceous community, then allowed trees to naturally colonize these communities (from intact seedbanks or as seed-rain) and grow. Nutrients and consumers had opposing effects on tree colonization and growth: adding nutrients and excluding consumers reduced tree colonization, but later increased established tree growth (height, basal diameter). Together, this shows stage-specific impacts of nutrients and consumers that may improve predictions of the rate and trajectory of succession: factors that initially limited tree colonization later helped established trees to grow.