Effects of density and ontogeny on size and growth ranks of three competing tree species

1. Rank reversals in species performance are theoretically important for structuring communities, maintaining diversity and determining the course of forest succession. Species growth ranks can change with ontogeny or in different microenvironments, but interactions between ontogeny and the environment are not well-understood because of the lack of long-term forest competition studies. While early differences in growth among species may reflect intrinsic differences in shade-tolerance and physiology, ontogenetic trends in growth and variation in neighbourhood density and composition may change or even reverse early patterns of growth rankings. 2. We experimentally studied spatial and temporal patterns of species interactions and growth for three northern tree species: Larix laricina, Picea mariana and Pinus strobus. We compared species size and growth rankings over an 11-year period, for different species mixtures planted at four density levels in north-eastern Minnesota, USA. 3. The benefits of different growth strategies changed with ontogeny and density leading to reversals in the size rank of competing species over time and space. High-density stands promoted dominance and resource pre-emption by L. laricina, whereas lower-density stands favoured gradual accumulation of biomass and eventual dominance by P. strobus. In the absence of strong neighbour competition, ontogenetic trends in growth had greater influence on growth patterns. 4. Species interactions affected the productivity of mixed stands vs. monocultures. Species generally grew more in monoculture than when planted with P. strobus at low density, or with L. laricina at high density. Only L. laricina and P. mariana showed potential for greater overall productivity, or over-yielding, when planted together than alone, probably because of improved resource uptake by the highly stratified canopy. 5. Synthesis. Density predictably determined whether size-asymmetric growth or ontogenetic growth trends would drive early establishment and growth patterns. Variation in vertical and horizontal structure that results from early competitive dynamics can influence the successional trajectory or character of the mature forest. This study extends previous efforts to identify the causes of rank reversals in communities and understand the importance of temporal changes beyond the early years of seedling establishment.