Grady, John M.; Quentin D. Read; Sydne Record; Nadja Ruegeer; Phoebe L. Zarnetske; Anthony I. Dell; Stephen P. Hubbell; Seat T. Michaletz and Brian J. Enquist

1. Both tree size and life history variation drive forest structure and dynamics, but little is known about how life history frequency changes with size. We used a scaling framework to quantify ontogenetic size variation and assessed patterns of abundance, richness, productivity and light interception across life history strategies from >114,000 trees in a primary, neotropical forest. We classified trees along two life history axes: a fast-slow axis characterized by a growth-survival trade -off, and a stature-recruitment axis with tall, long-lived pioneers at one end and short, short-lived recruiters at the other.2. Relative abundance, richness, productivity and light interception follow an approximate power law, systematically shifting over an order of magnitude with tree size. Slow saplings dominate the understorey, but slow trees decline to parity with rapidly growing fast and long-lived pioneer species in the canopy.3. Like the community as a whole, slow species are the closest to obeying the energy equivalence rule (EER)-with equal productivity per size class-but other life histories strongly increase productivity with tree size. Productivity is fuelled by resources, and the scaling of light interception corresponds to the scaling of productivity across life history strategies, with slow and all species near solar energy equivalence. This points towards a resource -use corollary to the EER: the resource equivalence rule.4. Fitness trade -offs associated with tree size and life history may promote coexistence in tropical forests by limiting niche overlap and reducing fitness differences.5. Synthesis. Tree life history strategies describe the different ways trees grow, survive and recruit in the understorey. We show that the proportion of trees with a pioneer life history strategy increases steadily with tree size, as pioneers become relatively more abundant, productive, diverse and capture more resources towards the canopy. Fitness tradeoffs associated with size and life history strategy offer a mechanism for coexistence in tropical forests.