Reports of drought-associated forest mortality have increased around the world, but the mechanisms of mortality are rarely direct in nature. Biotic agents may kill trees that could otherwise recover and can perpetuate and expand mortality after the stress is relieved. Sudden aspen decline (SAD) has caused rapid, widespread branch dieback and mortality of quak- ing aspen (Populus tremuloides Michx.). We compared insects and diseases in 162 damaged and neighboring healthy plots to determine contributing factors and their ecological roles. Cytospora canker, bronze poplar borer, and aspen bark beetles were the most common agents in damaged plots and correlated with crown loss and other factors related to SAD. This was the first documented outbreak of Trypophloeus populi, an aspen bark beetle. As bark beetles and bronze poplar borer in- creased in damaged stands, they tended to attack trees with healthier crowns. Environmental stress may have directly af- fected the success of these agents by increasing host susceptibility followed by a density-dependent increase in the insects' invasive ability. In contrast, Cytospora canker had an identical relationship to crown loss in healthy versus damaged plots, suggesting that it was not limited by inoculum but responded to host susceptibility. Most other pathogens and insects con- tributed little to SAD and appear to be primary or weakening agents. The biotic agents of mortality in a decline differ greatly from primary agents and play complex and varied roles in healthy versus declining stands.

Sudden aspen decline (SAD), affecting Populus tremuloides, was first observed in Colorado in 2004. By 2008 it affected at least 220,000ha, an estimated 17% of the aspen cover type in the state. In southwestern Colorado, we examined site and stand features in paired healthy and damaged plots to assess the effects of SAD on aspen and to identify factors associated with decline. Root mortality increased significantly with recent crown loss. Consequently, density of regeneration did not increase as the overstory deteriorated, and regeneration that originated since 2002 decreased significantly in stands with moderate to severe SAD. However, mortality of regeneration did not increase with that of the overstory. Remeasurement of a subset of plots after 1–2 yrs showed significant increases in severity. Contrary to expectations, overstory age and diameter were not related to SAD severity as measured by recent crown loss or mortality. Severity of SAD was inversely, but weakly, related to basal area, stem slenderness, and site index, and positively related to upper slope positions. This is consistent with moisture stress as an underlying factor. To test the role of climate as an inciting factor for SAD, a landscape-scale climate model was used to compare moisture status of declining and healthy aspen at the height of the warm drought in water year 2002. Polygons identified as damaged aspen in the 2008 aerial survey had greater moisture deficits than healthy aspen in the 2002 water year. SAD has led to loss of aspen cover in some stands, and is occurring in areas where early loss of aspen due to climate change has been predicted. Further warm, dry growing seasons will likely lead to recurrence of SAD.

Concentrated patches of recent trembling aspen (Populus tremuloides) mortality covered 515,091 ha of Colorado forests in 2006. Mortality has progressed rapidly. Area affected increased 58% between 2005 and 2006 on the Mancos-Dolores Ranger District, San Juan National Forest, where it equaled nearly 10% of the aspen cover type. In four stands that were measured twice, incidence of mortality increased from 7-9% in 2002/2003 to 31-60% in 2006. Mortality generally decreased with increasing elevation over the primary elevation range of aspen and occurred on less steep slopes than healthy aspen. Slope-weighted mean aspects of aspen cover type were northern at low elevations and generally southern at high elevations. Relative frequency of mortality was generally highest on southern to western aspects. In 31 stands measured in detail, mortality ranged from 0 to 100% (mean 32%) and was negatively correlated with stand density (P = 0.033). Size of trees affected was strongly correlated with amount of current mortality (P < 0.001), and current mortality was skewed toward larger diameter classes. Density of regeneration was in a low range typical of undisturbed stands and did not increase with overstory mortality.Agents that typically kill mature trees in aspen stands were unimportant in this mortality. Instead, a group of interchangeable, usually secondary agents was most commonly associated with mortality, including Cytospora canker (usually caused by Valsa sordida), aspen bark beetles (Trypophloeus populi and Procryphalus mucronatus), poplar borer (Saperda calcarata), and bronze poplar borer (Agrilus liragus). The rapidity of mortality, mortality agents involved, and probably other causal factors distinguish this phenomenon from the long-term loss of aspen cover usually attributed to successional processes operating in an altered disturbance regime (and often exacerbated by ungulate browsing). Our data are consistent with a hypothesis that (a) predisposing factors include stand maturation, low density, southern aspects and low elevations; (b) a major inciting factor was the recent, acute drought accompanied by high temperatures, and; (c) contributing factors and proximate agents of mortality are the common biotic agents observed. On sites with poor regeneration and weak root systems, clones may die, resulting in the long-term loss of aspen forest cover. Published by Elsevier B.V.