Natural disturbance and gap phase dynamics in southern Appalachian spruce–fir forests

Abstract
Spatially small canopy gaps dominated the natural disturbance regime of old-growth spruce–fir forests in the Great Smoky Mountains, North Carolina and Tennessee. New gaps ≤200 m2 in size were formed with a frequency of 0.006 to 0.009/year and the 1- to 10-year age class of these gaps covered an estimated 6 to 17% of the study area (depending on calculation method). Sixty canopy gaps sampled on a 7-ha intensive study site ranged in size from 15 to 150 m2. Although tree replacement patterns in these gaps were unpredictable from gap size and age, the gap event was important in species interactions. The three canopy dominants, Abiesfraseri (Pursh) Poir., Picearubens Sarg., and Betulalutea Michx.f., had unique suites of life history traits. Abies reached high understory densities, but had the highest canopy turnover rate of the three species. Betula was scarce in the understory, but had a crown expansion rate in gaps eight times that of the two conifers. Picea was the longest lived species and appeared to have the best survivorship. Tree replacement models based on advanced regeneration led to the prediction that Abies canopy density would increase and Betula canopy density would decrease, while models based on gap inventories led to opposite conclusions. Data from direct observation of the gap capture process supported the hypothesis that this old-growth stand was near compositional equilibrium and underscored the importance of disturbance effects in community organization.