Death by attrition: a hypothesis for wave mortality of subalpine Abiesbalsamea

Abstract
Wave mortality in subalpine Abies forests of the northeastern United States is characterized by the progressive dieback of overstory dominants in a compact zone that advances systematically through mature stands and is followed by vigorous regeneration of the overstory species. The principal driving force for this dieback appears to be exposure at the edge of canopy gaps to very high winds. In this study, individual trees at the leading edge of a dieback front and trees in a thinned plot within the mature stand downwind of the advancing wave both experienced a substantial decline in foliage mass and, consequently, in prebudbreak foliar starch pool over a 2-year period. In the case of the thinned stand, this reduction was attributed to a large increase in wintertime green litter fall. Increased exposure at the wave front and in the thinned stand also resulted in a dramatic increase in root damage relative to control trees. Coincidental to breakage of roots, root xylem discoloration (attributed to wounding and invasion by root pathogens) was greater on trees at the wave front than on trees in the mature zone. Loss of active foliage and roots was accompanied by a two- to five-fold reduction in annual wood increment among dieback zone trees. These data suggest that exposed trees, which suffer a substantial reduction in crown size and foliage carbohydrate pool in an environment where growth is marginal to begin with, may not be able to keep up with the increased energy demand for root and foliage turnover and eventually suffer death by attrition of both tissues.