Neighborhood competition mediates crown development of Picea sitchensis in Olympic rainforests: Implications for restoration management

Abstract

Large trees are critically important for structuring ecosystems and providing habitat, and trees with complex crowns provide more of these services than comparably sized trees with simple crowns. Forest managers are increasingly emulating old-growth structure by retaining various densities of aggregated and dispersed trees. This study explores how individual within-crown complexity develops in a wide range of tree densities and interprets them in terms of management goals. Stratified LiDAR data were used to select 36 Picea sitchensis trees 55 to 91 m tall and from the complete range of tree densities in Olympic rainforests. These were climbed, three-dimensionally mapped—including the trunk and every appendage—to quantify tree allometry and habitat-related structures within their crowns. Sampled trees were then aged with crossdated increment cores to construct a chronosequence of crown development. The sample ranged from 70 to 250 cm diameter and 100 to 400 years old. Circular 30 m radius plots were installed around each tree to quantify neighborhood density. An additional 19 trees ranging from 10 to 94 m tall were added to the sample (N = 55) to describe baseline crown development and then compared to the 36-tree sample in which we modeled how this baseline is altered by neighborhood competition. After ∼200 yr, crown mass increases faster than trunk mass due to accumulation and growth of large appendages. The largest neighbors within 25 m compete with Picea, reducing trunk and appendage diameters as well as crown volume by roughly 20% for each 25% increase in density for a given tree age. Reduced growth and higher mortality of lower-crown appendages in dense forests delays development of large (>15 cm diameter) appendages for over 100 yr. Abundance of structures associated with reiterated trunks is unrelated to neighborhood density but increases with crown damage. These findings contribute to an empirically-based conceptual framework for crown development in Picea and interpretation of management guidelines. Picea is then compared to Pseudotsuga menziesii—another well-studied tall conifer of Olympic rainforests. Picea accumulates ecologically important crown structures much faster than Pseudotsuga yet has roughly half the longevity, indicating that these co-occurring species have complementary functions. Because of its shade tolerance and extremely rapid growth in rainforests, Picea can be used to accelerate development of complex crown structures in forests typically planted with Pseudotsuga after logging.