Abstract

Despite its importance to biodiversity and ecosystem function, patterns and drivers of regional scale variation in forest structure and development are poorly understood. We characterize structural variation, create a hierarchical classification of forest structure, and develop an empirically based framework for conceptualizing structural development from 11,091 plots across 25 million ha of all ownerships in Oregon, Washington, and inland Northern California, USA. A single component related to live tree biomass accounted for almost half of the variation in a principal components analysis of structural attributes, but components related to live tree density and size, dead wood, and understory vegetation together accounted for as much additional variation. These results indicate that structural development is more complex than a monotonic accumulation of live biomass as other components may act independently or emerge at multiple points during development. The classification revealed the diversity of structural conditions expressed at all levels of live biomass depending on the timing and relative importance of a variety of ecological processes (e.g., mortality) in different vegetation zones. Low live biomass structural types (<25 Mg/ha) illustrated the diversity of early‐seral conditions and differed primarily in density of live trees and abundance of snags and dead wood. Moderate live biomass structural types (25–99 Mg/ha) differed in tree size and density and generally lacked dead wood, but some structurally diverse types associated with partial stand‐replacing disturbance had abundant live and dead legacies. High live biomass structural types (>100 Mg/ha) substantiated the diversity of later developmental stages and exhibited considerable variation in the abundance of dead wood and density of big trees. Most structural types corresponded with previously described stages of development, but others associated with protracted early development, woodland/savannah transitions, and partial stand‐replacing disturbance lacked analogs and indicated alternative pathways of development. We propose a conceptual framework that distinguishes among families of pathways depending on the range of variation along different components of structure, the relative importance of different disturbances, and complexity of pathways. Our framework is a starting point for developing more comprehensive models of structural development that apply to a wider variety of vegetation zones differing in environment and disturbance regimes.

Highlights

  • Despite the importance of forest structure to ecosystem function (Waring and Running 2007) and biodiversity (MacArthur and MacArthur 1961), variation in structure at the regional scale is poorly understood

  • This study provides the first empirically based regional-scale characterization of forest structure and how it varies in relation to age, vegetation zone, and disturbance history

  • Most of variation in forest structure was accounted for by live tree biomass, other components related to tree size and density, dead wood, and understory vegetation cumulatively accounted for as much variation

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Summary

Introduction

Despite the importance of forest structure to ecosystem function (Waring and Running 2007) and biodiversity (MacArthur and MacArthur 1961), variation in structure at the regional scale is poorly understood. Past regional scale studies have focused primarily on how climate constrains biomass (Gholz 1982, Malhi et al 2006, Urquiza-Haas et al 2007, Hudiburg et al 2009), but more detailed investigations of other structural attributes such as standing dead trees (snags) and dead or downed wood are rare (Spies and Franklin 1991, Ohmann and Waddell 2002, Ohmann et al 2007, Ares et al 2012). Snag abundance is often quantified by density (per unit area), but because dead wood is continually breaking down through fragmentation and decomposition, biomass estimates offer more precise measures of the abundance of snags and dead and downed wood in ecosystem studies (Harmon et al 1986). Forest structure is an inherently complex multivariate concept with broad ecological implications that vary by individual components and ecological context

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